Pacific West Coast Research Articles

Whole genome sequencing of Canadian Saccharomyces cerevisiae strains isolated from spontaneous wine fermentations reveals a new Pacific West Coast Wine clade.

Vineyards in wine regions around the world are reservoirs of yeast with oenological potential. Saccharomyces cerevisiae ferments grape sugars to ethanol and generates flavor and aroma compounds in wine. Wineries place a high-value on identifying yeast native to their region to develop a region-specific wine program. Commercial wine strains are genetically very similar due to a population bottleneck and in-breeding compared to the diversity of S. cerevisiae from the wild and other industrial processes. We have isolated and microsatellite-typed hundreds of S. cerevisiae strains from spontaneous fermentations of grapes from the Okanagan Valley wine region in British Columbia, Canada. We chose 75 S. cerevisiae strains, based on our microsatellite clustering data, for whole genome sequencing using Illumina paired-end reads. Phylogenetic analysis shows that British Columbian S. cerevisiae strains cluster into 4 clades: Wine/European, Transpacific Oak, Beer 1/Mixed Origin, and a new clade that we have designated as Pacific West Coast Wine. The Pacific West Coast Wine clade has high nucleotide diversity and shares genomic characteristics with wild North American oak strains but also has gene flow from Wine/European and Ecuadorian clades. We analyzed gene copy number variations to find evidence of domestication and found that strains in the Wine/European and Pacific West Coast Wine clades have gene copy number variation reflective of adaptations to the wine-making environment. The "wine circle/Region B", a cluster of 5 genes acquired by horizontal gene transfer into the genome of commercial wine strains is also present in the majority of the British Columbian strains in the Wine/European clade but in a minority of the Pacific West Coast Wine clade strains. Previous studies have shown that S. cerevisiae strains isolated from Mediterranean Oak trees may be the living ancestors of European wine yeast strains. This study is the first to isolate S. cerevisiae strains with genetic similarity to nonvineyard North American Oak strains from spontaneous wine fermentations.

Genetic analyses reveal a non-panmictic genetic structure in the sablefishAnoplopoma fimbriain the northern Pacific

AbstractThe sablefish Anoplopoma fimbria is distributed on the continental shelf of the North Pacific, has a high commercial value for both fisheries and aquaculture, and represents a shared resource between countries in the North Pacific basin. In the present study, we extend the geographic range surveyed in previous studies and reexamine the population’s genetic structure by integrating phylogeographic patterns of mitochondrial DNA and microsatellite DNA markers. Our results contradict the proposal that sablefish constitute a single population throughout their distributional range. We observed a signal of ancient divergence in mtDNA that differentiates the North Pacific West Coast from the rest of the sample sites, and microsatellite markers reveal a contemporary isolation of Mexican sablefish. Our findings show genetic differences between localities that should be explored in more detail to fully understand the interconnectedness that appears to exist between populations.

A chromosome-level reference genome for the giant pink sea star, Pisaster brevispinus, a species severely impacted by wasting.

Efforts to protect the ecologically and economically significant California Current Ecosystem from global change will greatly benefit from data about patterns of local adaptation and population connectivity. To facilitate that work, we present a reference-quality genome for the giant pink sea star, Pisaster brevispinus, a species of ecological importance along the Pacific west coast of North America that has been heavily impacted by environmental change and disease. We used Pacific Biosciences HiFi long sequencing reads and Dovetail Omni-C proximity reads to generate a highly contiguous genome assembly of 550 Mb in length. The assembly contains 127 scaffolds with a contig N50 of 4.6 Mb and a scaffold N50 of 21.4 Mb; the BUSCO completeness score is 98.70%. The P. brevispinus genome assembly is comparable to the genome of the congener species P. ochraceus in size and completeness. Both Pisaster assemblies are consistent with previously published karyotyping results showing sea star genomes are organized into 22 autosomes. The reference genome for P. brevispinus is an important first step toward the goal of producing a comprehensive, population genomics view of ecological and evolutionary processes along the California coast. This resource will help scientists, managers, and policy makers in their task of understanding and protecting critical coastal regions from the impacts of global change.

Of Martyrs and “Social Dynamites”: The Ghadar and IWW in California

ABSTRACT In the early decades of the twentieth century, the Pacific West Coast of the USA gradually turned into a hotbed of Indian anti-colonial activity. California, in particular, became a point of convergence for Bengali radicals/intellectuals and Punjabi migrant workers. The Ghadar would gradually emerge in 1913 out of the interactions between these two elements. My paper is animated by the understanding that zooming in on Indian anti-colonial activity can be a very productive way of situating California in a transnational historical framework. In my paper, I focus on the interface between the Ghadar movement and the IWW in California. Existing studies have focused on superficial similarities between the two radical movements as a way of explaining their mutual compatibility. Departing from such approaches, I will try to show that similar understandings of time, force, and rupture rendered the Ghadar and IWW compatible at a very fundamental level.

Evaluating Chinook salmon (Oncorhynchus tshawytscha) response to artificial light in support of bycatch mitigation

In commercial trawl fisheries in the North Pacific and US West Coast, fishermen and scientists are evaluating if artificial lights facilitate escapement of bycaught Chinook salmon (Oncorhynchus tshawytscha) from the trawl by attracting them to an opening provided by a bycatch reduction device. Inconsistent behaviour and escapement rates when lights were used in the trawl led us to conduct a laboratory study to evaluate the role of light properties (intensity, colour, and strobe) on marine Chinook salmon behaviour. Results from this study suggest a negative phototactic response. Light colour and strobe, and the interaction between them, differentially affected behavioural response with regard to mean swimming speed and distance from and habituation to the light. White light intensity had limited influence on response; however, the range of trialed intensities was limited. While behaviour is contextual and responses in a laboratory setting cannot be directly extrapolated to responses in fishing gear, this study highlights the significant role of light properties when trying to affect behaviour for bycatch mitigation and the importance of distinguishing between a response to light and to illuminated surroundings.

Ommastrephid squid paralarvae potential nursery habitat in the tropical-subtropical convergence off Mexico

The Gulf of California and adjacent Pacific have passed through a prolonged warming period that affects the Humboldt squid D. gigas life cycle, which was the third largest fishery in Mexico until its total collapse after 2010. The aims of the present study were (i) to characterize the nursery habitat of paralarvae of the complex Sthenoteuthis oualaniensis–Dosidicus gigas (SD complex) on the basis of 1112 paralarvae collected from zooplankton trawls carried out during ten oceanographic cruises off northwestern Mexico between 2010 and 2017; and (ii) to model the seasonal and interannual variability of the nursery habitat, applying a generalized additive model (GAM) using a regional long-term (1922–2017) hydrographic database. The GAM showed significant relationships of paralarvae abundance with the seasonal average of thermocline depth, absolute salinity, and conservative temperature. The potential nursery habitat of the SD complex paralarvae showed its maximum expected abundance in the entrance of the Gulf of California during spring, when the thermocline depth was shallowest (<30 m). During summer the nursery habitat moved northward along the Pacific west coast of the peninsula. Autumn and winter were less suitable seasons for paralarvae nursery because the thermocline was deep throughout the area of study. Potential nursery habitat did not change interannually in location during La Niña conditions, but did increase the expected abundance in its southern extension. El Niño events contracted the nursery habitat near Cabo San Lucas, and caused the habitat to move northwards over both coasts of the Baja California peninsula. Despite seasonal and interannual climatic variability, two regions remained suitable habitats almost throughout the year. One was associated with coastal upwelling regions (Cabo Corrientes) and the second comprised the oceanic region of high mesoscale kinetic energy located south of Cabo San Lucas. We conclude that the potential nursery habitat of the SD complex is more coastal than oceanic, and is associated with a shallow thermocline in the tropical-subtropical convergence off Mexico. The recent anomalous warming period in this region could drive the changes in the morphology of D. gigas in the Gulf of California, given the effects of environmental variability over early stages of development throughout the adult ontogenesis.

Signals of resilience and change in tidepool fish communities on the Pacific coast of Vancouver Island, Canada

AbstractAimOcean warming and marine heatwaves have emerged as strong predictors of biodiversity change in coastal systems. Here, we test for systematic change in tidepool fish communities between recent and historical data sets, which include the North Pacific marine heatwave of 2014–2016.LocationPeriodic sampling occurred since 1966 at study site isolated from human activities on the Pacific west coast of Vancouver Island, Canada.MethodsRepeated fish sampling at seven wave‐exposed rocky shore tidepools was conducted on 8 (historical: 1966–1993) and 5 (recent: 2012–2016) occasions during low tide using methods that capture all individuals present (including cryptic and rare species). Species were subsequently identified using standard taxonomic approaches and abundances recorded. These data comprised the basis for analyses of both community‐level and species‐specific responses to test for differences in the historical and recent time periods.ResultsWe did not detect systematic increases or declines in species density, total abundance of fishes within tidepools and community similarity between historical and recent sampling events in high‐intertidal pools. However, fish communities from low‐shore pools decreased in the number of species present, overall abundance and community similarity. Those species lost from the lower pools were non‐resident rare species typical of subtidal environments. By contrast, a warm‐affinity crevice kelpfish was observed only recently. We also found that six species had higher or lower abundances between historical and recent sampling events, and one species occurred at higher vertical heights.Main conclusionFish communities in high‐intertidal pools were resilient between historical and recent sampling periods, and lacked evidence of invasion. However, some signals of community and species changes were evident in the lower shore. Overall, these communities may reach a threshold with ongoing ocean warming and continued monitoring is required to reveal any major changes as they unfold.

An Arts Organization Policy Brief on Indigenous Knowledges

This policy brief provides an overview of Indigenous knowledges for an arts organization on the Pacific West Coast in Canada. To orient readers, the brief is contextualized within the broader arc of the organizations' history of commitments to, departures from, and re-engagement with commitments to decolonization and decolonial practice. It provides a list of additional resources.

Correction to: Efficiency drivers for the South Pacific West coast port terminals: a two-stage non-convex metafrontier dea approach

Correction to: Efficiency drivers for the South Pacific West coast port terminals: a two-stage non-convex metafrontier dea approach

Efficiency drivers for the South Pacific West coast port terminals: a two-stage non-convex metafrontier dea approach

We measure technical efficiency of Peruvian and Chilean port terminals, to evaluate the influence of certain contextual variables in the terminals’ efficiency levels. The sample includes 14 port terminals from 2004 to 2014. Due to the potential differences, we have estimated a DEA model in a non-convex metafrontier framework. Afterwards, we estimated all the regression models proposed in the literature that could be used to explain not only the technical efficiency estimated with respect to the metafrontier (TE*) but also each one of its components: the technical efficiency with respect to the group-specific frontier (TEk) and the technological gap ratio (TGR). Results are robust across models.

Understanding Color Polymorphism in the Sea Star, Pisaster ochraceus, Through Activity Levels and Dietary Preference

Color polymorphism is found in a wide array of organisms ranging from copepods to black bears. Pisaster ochraceus, an intertidal sea star on the North Pacific West Coast, shows a striking color polymorphism including a range of orange, brown, maroon, and purple. Pisaster shows extensive geographic color variation, with &gt;95% frequency of bright purple Pisaster in the Vancouver region; while the open pacific coast has a frequency distribution of 6-28% orange, 68-90% reddish-brown to dull purple, and a small percentage of bright purple. Maintenance of color polymorphism remains unknown, but one hypothesis suggests an underlying genetic component with regional-scale variation controlled by an ecological factor. Two ecological factors suggested are diet and salinity. This study analyzes the differences in dietary preference and activity levels between the color morphs in Bamfield, British Columbia. Pisaster was binned into two colors, orange and purple, for statistical analysis. Feeding laboratory trials showed no preference between the color morphs for Mytilus edilus or Mytilus californianus. Self-righting trials were performed in a low (20 psu) and controlled (35 psu) salinity and no significant difference was found between the color morphs. Interestingly, the activity levels of orange color morphs were not significantly different between low and control salinity. The results suggest there are ecological differences between the color morphs that still need to be evaluated.

Tsunami detection by high-frequency radar in British Columbia: performance assessment of the time-correlation algorithm for synthetic and real events

The authors recently proposed a new method for detecting tsunamis using high-frequency (HF) radar observations, referred to as “time-correlation algorithm” (TCA; Grilli et al. Pure Appl Geophys 173(12):3895–3934, 2016a, 174(1): 3003–3028, 2017). Unlike standard algorithms that detect surface current patterns, the TCA is based on analyzing space-time correlations of radar signal time series in pairs of radar cells, which does not require inverting radial surface currents. This was done by calculating a contrast function, which quantifies the change in pattern of the mean correlation between pairs of neighboring cells upon tsunami arrival, with respect to a reference correlation computed in the recent past. In earlier work, the TCA was successfully validated based on realistic numerical simulations of both the radar signal and tsunami wave trains. Here, this algorithm is adapted to apply to actual data from a HF radar installed in Tofino, BC, for three test cases: (1) a simulated far-field tsunami generated in the Semidi Subduction Zone in the Aleutian Arc; (2) a simulated near-field tsunami from a submarine mass failure on the continental slope off of Tofino; and (3) an event believed to be a meteotsunami, which occurred on October 14th, 2016, off of the Pacific West Coast and was measured by the radar. In the first two cases, the synthetic tsunami signal is superimposed onto the radar signal by way of a current memory term; in the third case, the tsunami signature is present within the radar data. In light of these test cases, we develop a detection methodology based on the TCA, using a correlation contrast function, and show that in all three cases the algorithm is able to trigger a timely early warning.

Subregional Comparison for Forest- to-Product Biomass Supply Chains on the Pacific West Coast, USA

Abstract. Transportable biomass conversion facilities producing biochar, briquettes, and torrefied wood are modeled and optimized for five different sub-regions within the Pacific Northwest. Subregional case studies in Quincy, California; Lakeview, Oregon; Oakridge, Oregon; Port Angeles, Washington; and Warm Springs, Oregon, are evaluated to characterize the potential economic viability of these novel transportable designs. A mixed integer program is used to characterize the supply chain from residue extraction to market optimizing transportation, production, and plant mobility in order to minimize the supply chain costs. Regional variations including log specifications, energy rates, trucking, and logistic capacities are considered within the model and supporting analyses to differentiate regional costs and market viabilities. It was found that the optimal transportable design included facility movement on a 1 to 2.5 year frequency depending on product and region with biochar being the most likely to be economically viable. Regional feedstock composition and availability was the biggest indicator of lower cost production. Supply chain costs varied by 5%-10% depending on product and region being produced. Transportation and mobilization were found to account for 15%-30% of the overall supply chain cost. Quincy, California, and torrefied wood were found to have the lowest of these costs due to low move frequency and high wood availability while Port Angeles, Washington, with briquettes was the highest. With regards to fuel price sensitivity, torrefied wood was the most sensitive as its conversion process was most energy intensive (±12%-13%) and biochar least sensitive (±3-5%).Transportation accounted for 5% to 30% of the fuel price variation due to diesel prices depending on product and region. When including grid-connectivity, cost reductions were approximately 6%-7% for biochar, 27%-29% for briquettes and 33%-38% for torrefied wood. These findings indicate biochar as the most likely candidate for a transportable conversion system given its relatively low power consumption, high allowable moisture content, and low product transportation cost. Quincy, California, was found to be the most desirable sub region with the lowest overall production costs attributed to its high input quality feedstock and relative accessibility; its higher grid-connected power cost also makes transportable options relatively more attractive than other off-grid locations. Port Angeles, WA had the highest production costs and lowest grid-energy costs. Our results indicate that a rise in diesel price, while incentivizing transportable conversion facilities due to more cost effective transportation, would be more than offset by the higher cost energy consumption during the conversion process when compared with grid-power with the potential exception of biochar. Overall, we see a transportable operation with grid-power would likely be the difference between an economically viable supply chain and one that is not. Keywords: Biomass supply, Biomass products, Facility location, Mixed integer programming, Strategic planning, Transportable plants.

Restoring lepidopteran diversity in a tropical dry forest: relative importance of restoration treatment, tree identity and predator pressure.

Tropical dry forests (TDFs) have been widely transformed by human activities worldwide and the ecosystem services they provide are diminishing. There has been an urgent call for conservation and restoration of the degraded lands previously occupied by TDFs. Restoration experiences aim to recover species diversity and ecological functions. Different restoration strategies have been used to maximize plant performance including weeding, planting or using artificial mulching. In this investigation, we evaluated whether different restoration practices influence animal arrival and the reestablishment of biotic interactions. We particularly evaluated lepidopteran larvae diversity and caterpillar predation on plants established under different restoration treatments (mulching, weeding and control) in the Pacific West Coast of México. This study corroborated the importance of plant host identity for lepidopteran presence in a particular area. Lepidopteran diversity and herbivory rates were not affected by the restoration treatment but they were related to tree species. In contrast, caterpillar predation marks were affected by restoration treatment, with a greater number of predation marks in control plots, while caterpillar predation marks among plant species were not significantly different. This study highlights the importance of considering the introduction of high plant species diversity when planning TDF restoration to maximize lepidopteran diversity and ecosystem functioning.

Different soil moisture control of net methane oxidation and production in organic upland and wet forest soils of the Pacific coastal rainforest in Canada

In a changing climate, understanding how soil hydrology impacts greenhouse gas dynamics will be important for the future management of the soils in the forests on the Canadian Pacific west coast. In a laboratory study, the impact of soil hydrology on potential net methane (CH4) exchange rates and the abundance of methanotrophs (CH4oxidation) and methanogens (CH4production) in upland and water-saturated wet soils were investigated. CH4oxidation and production rates were highest in the wet soils, which corresponded to higher numbers of methanotrophs and methanogens, indicating a link between the microbial abundance and CH4exchange rates. Also, CH4production was induced in the upland soils, indicating the presence of methanogens. The optimum soil moisture content for CH4oxidation was highest in upland soils and the wet soils sustained higher CH4oxidation rates over a broader range of soil moisture. These results underline the importance of the soil hydrological controls of CH4oxidation in contrasting soils and forest types, which deserves further attention in field-based studies.

Redescription of Dynoides elegans (Boone, 1923) (Crustacea, Isopoda, Sphaeromatidae) from the north-eastern Pacific.

Dynoides elegans (Boone, 1923) from southern California is reviewed, redescribed, and figured. The original species description did not include figures, making it difficult to attribute individuals to the species. Dynoides saldanai Carvacho and Haasmann, 1984 and Dynoides crenulatus Carvacho & Haasman, 1984 from the Pacific Coast of Mexico and Dynoides brevicornis Kussakin & Malyutina, 1987, from Furugelm Island, Peter the Great Gulf in the Sea of Japan, appear morphologically more similar to each other than to western Pacific species. A large pleonal process is present in about half of the Dynoides species, but is absent in this north-eastern Pacific clade and the north-western Pacific Dynoides brevicornis and Dynoides brevispina. Dynoides dentisinus Shen, 1929 possess a large pleonal spine. It is known from China, Japan, and Korea and is introduced in San Francisco Bay; it can be easily distinguished from Dynoides elegans by the presence of a pleonal process in the former. A key to the Pacific West Coast Dynoides is provided.

Status of Exosphaeromaamplicauda (Stimpson, 1857), E.aphrodita (Boone, 1923) and description of three new species (Crustacea, Isopoda, Sphaeromatidae) from the north-eastern Pacific.

Exosphaeroma amplicauda (Stimpson, 1857) from the west coast of North America is reviewed and redescribed and revealed to be a group of closely related species. A neotype is designated and the species redescribed based on the neotype and topotypic specimens. Exosphaeroma amplicauda is known only from the coast of California, at Marin, Sonoma and San Mateo Counties. Exosphaeroma aphrodita (Boone, 1923), type locality La Jolla, California and previously considered nomen dubium is taken out of synonymy and re-validated. A further three species: Exosphaeroma paydenae sp. n., Exosphaeroma russellhansoni sp. n., and Exosphaeroma pentcheffi sp. n. are described herein. Sphaeroma octonctum Richardson, 1899 is placed into junior synonymy with Exosphaeroma amplicauda. A key to the Pacific West Coast Exosphaeroma is provided.

Bacterial Populations (First Record) at Two Shallow Hydrothermal Vents of the Mexican Pacific West Coast

Thermophilic and metal-oxidizing bacteria were identified in shallow hydrothermal vents on the western Mexican coast. The role of these bacteria in biomineralization processes observed in the vents is explained, and the effect of the vents on biodiversity of prokaryotes is discussed. Research was done at two shallow hydrothermal vent sites: Bahía Concepción (BC) in the Baja California Peninsula and Punta Mita (PM), on the central Pacific coast. Temperature at the sediments proximal to the vents was similar, but the redox potentials (0.5 V in BC and −0.3 V in PM) and pH (6.2 in BC and as low as 4.5 in PM) differed. The composition of the discharged water ranged from nearly seawater to lower-salinity fluids, and the gas phase was mainly CO2 at BC and N2 and CH4 at PM. The study focuses on the biogeochemical processes related to the different species of bacteria present in the studied sites, which are involved in the anaerobic oxidation of methane (AOM), seawater sulfate reduction, and metal oxidation. The detected bacterial lineages represented typical deep vent species, which disproves a previous hypothesis that proposed that different consortia were populating deep and shallow hydrothermal vents. The results obtained here show that the main parameter affecting the bacterial groups present in shallow vents was the redox potential: gamma-, delta-, and epsilon proteobacteria as well as Bacteriodetes are present under the oxidizing conditions of BC, and Thermotogae, Aquificae, and Planctomycetes are present in PM. Sunlight abundance favored the prevalence of halophilic and Chlorofleaxae bacteria in both areas.

Phenotypic plasticity of invasive Spartina densiflora (Poaceae) along a broad latitudinal gradient on the Pacific Coast of North America

Phenotypic acclimation of individual plants and genetic differentiation by natural selection within invasive populations are two potential mechanisms that may confer fitness advantages and allow plants to cope with environmental variation. The invasion of Spartina densiflora across a wide latitudinal gradient from California (USA) to British Columbia (Canada) provides a natural model system to study the potential mechanisms underlying the response of invasive populations to substantial variation in climate and other environmental variables. We examined morphological and physiological leaf traits of Spartina densiflora plants in populations from invaded estuarine sites across broad latitudinal and climate gradients along the Pacific west coast of North America and in favorable conditions in a common garden experiment. Our results show that key foliar traits varied widely among populations. Most foliar traits measured in the field were lower than would be expected under ideal growing conditions. Photosynthetic pigment concentrations at higher latitudes were lower than those observed at lower latitudes. Greater leaf rolling, reduced leaf lengths, and lower chlorophyll and higher carbon concentrations were observed with anoxic sediments. Lower chlorophyll to carotenoids ratios and reduced nitrogen concentrations were correlated with sediment salinity. Our results suggest that the variations of foliar traits recorded in the field are a plastic phenotypic response that was not sustained under common garden conditions. SPARTINA DENSIFLORA shows wide differences in its foliar traits in response to environmental heterogeneity in salt marshes, which appears to be the result of phenotypic plasticity rather than genetic differentiation.

Components of ecosystem respiration and an estimate of net primary productivity of an intermediate-aged Douglas-fir stand

Continuous half-hourly chamber-measured belowground ( R s) and eddy covariance (EC)-measured total ecosystem ( R e) respiration in a 56-year-old Pacific west coast Douglas-fir stand from 2003 to 2005 were analyzed to study their seasonal and interannual variability. Seasonal variation in both R s and R e was better predicted by soil temperature at the 5 cm depth than at any other depth and air temperature. R e was more responsive than R s to changes in temperature and soil water content. While nighttime ( R sn) and daytime ( R sd) soil respiration showed no difference in their response to soil temperature, daytime ecosystem respiration ( R ed) was less responsive than its nighttime counterpart ( R en) to changes in soil temperature. Half-hourly R ed was almost always smaller than R en possibly due to photoinhibition while daily total R ed was higher than daily total R en during summer and spring months but smaller in winter and autumn months with the latter due to fewer daylight hours. A distinct seasonal pattern in the R s to R e ratio was observed. The 3-year mean seasonal R s/ R e was at its minimum of 0.52 in spring followed by 0.63 in summer, 0.81 in autumn and a maximum of 0.86 in winter. Both daily total and half-hourly R sd/ R ed were larger than R sn/ R en. It appears that the different responses of R s and R e to environmental variables arise as a result of seasonal variations in photosynthesis, mobilization and use of stored carbohydrates, and differences in the phenology of aboveground and belowground plant tissues. On an annual basis, R s accounted for 62% of R e with the latter accounting for 86% of the carbon (C) assimilated in annual photosynthesis or gross primary productivity (GPP), leaving the net C sequestration efficiency (1 − R e/GPP) at 14%. In the relatively dry year of 2003, both R s and R e as well as GPP were the lowest but with the highest net ecosystem productivity (NEP) of the 3 years. The relatively wetter growing season soil moisture regime in 2005 resulted in the highest R e and GPP. Partitioning of R s into its autotrophic and heterotrophic components indicated that 54% of GPP was respired back to the atmosphere as autotrophic respiration and an additional 32% was lost in the decomposition of litterfall and soil organic matter. The mean annual estimate of net primary productivity (NPP) at 843 g C m −2 accounted for 47% of the mean annual GPP of 1815 g C m −2.