Sea urchins in a high CO2 world: partitioned effects of body-size, ocean warming and acidification on metabolic rate

Body-size and temperature are the major factors explaining metabolic rate, and the additional factor of pH is a major driver at the biochemical level. These three factors have frequently been found to interact, complicating the formulation of broad models predicting metabolic rates and hence ecological functioning. In this first study of the effects of warming and ocean acidification, and their potential interaction, on metabolic rate across a broad body-size range (two-to-three orders of magnitude difference in body mass) we addressed the impact of climate change on the sea urchin Heliocidaris erythrogramma in context with climate projections for east Australia, an ocean warming hotspot. Urchins were gradually introduced to two temperatures (18 and 23 °C) and two pH (7.5 and 8.0), and maintained for two months. That a new physiological steady-state had been reached, otherwise know as acclimation, was validated through identical experimental trials separated by several weeks. The relationship between body-size, temperature and acidification on the metabolic rate of H. erythrogramma was strikingly stable. Both stressors caused increases in metabolic rate; 20% for temperature and 19% for pH. Combined effects were additive; a 44% increase in metabolism. Body-size had a highly stable relationship with metabolic rate regardless of temperature or pH. None of these diverse drivers of metabolism interacted or modulated the effects of the others, highlighting the partitioned nature of how each influences metabolic rate, and the importance of achieving a full acclimation state. Despite these increases in energetic demand there was very limited capacity for compensatory modulating of feeding rate; food consumption increased only in the very smallest specimens, and only in response to temperature, and not pH. Our data show that warming, acidification and body-size all substantially affect metabolism and are highly consistent and partitioned in their effects, and for H. erythrogramma near-future climate change will incur a substantial energetic cost.

Effects of pre-hatching pCO2 on development and survival of zoea stages of Arctic spider crab Hyas araneus

Sensitivity of marine crustaceans to anthropogenic CO2 emissions and the associated acidification of the oceans may be less than that of other, especially lower, invertebrates. However, effects on critical transition phases or carry-over effects between life stages have not comprehensively been explored. Here we report the impact of elevated seawater PCO2 values (3100 µatm) on Hyas araneus during the last 2 weeks of their embryonic development (pre-hatching phase) and during development while in the consecutive zoea I and zoea II larval stages (post-hatching phase). We measured oxygen consumption, dry weight, developmental time and mortality in zoea I to assess changes in performance. Feeding rates and survival under starvation were investigated at different temperatures to detect differences in thermal sensitivities of zoea I and zoea II larvae depending on pre-hatch history. When embryos were pre-exposed to elevated PCO2 during maternal care, mortality increased about 60% under continued CO2 exposure during the zoea I phase. The larvae that moulted into zoea II, displayed a developmental delay by about 20 days compared to larvae exposed to control PCO2 during embryonic and zoeal phases. Elevated PCO2 caused a reduction in zoea I dry weight and feeding rates, while survival of the starved larvae was not affected by the seawater CO2 concentration. In conclusion, CO2 effects on egg masses under maternal care carried over to the first larval stages of crustaceans and reduced their survival and development to levels below those previously reported in studies exclusively focussing on acute PCO2 effects on the larval stages.

Diversidad funcional y diversidad filogenética en los bosques secos del sur del Ecuador

El concepto tradicional de reglas de ensamblaje refleja la idea de que las especies no co-ocurren al azar sino que están restringidos en su co-ocurrencia por la competencia interespecífica o por un filtrado ambiental. En está tesis abordé la importancia de los procesos que determinan el ensamble de la comunidad en la estructuración de los Bosques Secos en el Sur del Ecuador. Este estudio se realizó en la región biogeográfica Tumbesina, donde se encuentra la mayor concentración de bosques secos tropicales bien conservados del sur de Ecuador, y que constituyen una de las áreas de endemismo más importantes del mundo. El clima se caracteriza por una estación seca que va desde mayo a diciembre y una estación lluviosa de enero a abril, su temperatura anual varía entre 20°C y 26°C y una precipitación promedio anual entre 300 y 700 mm. Mi primer tema fue orientado a evaluar si la distribución de los rasgos funcionales a nivel comunitario es compatible con la existencia de un filtro ambiental (filtrado del hábitat) o con la existencia de un proceso de limitación de la semejanza funcional impuesta por la competencia inter-específica entre 58 especies de plantas leñosas repartidas en 109 parcelas (10x50m). Para ello, se analizó la distribución de los valores de cinco rasgos funcionales (altura máxima, densidad de la madera, área foliar específica, tamaño de la hoja y de masa de la semilla), resumida mediante varios estadísticos (rango, varianza, kurtosis y la desviación estándar de la distribución de distancias funcionales a la especies más próxima) y se comparó con la distribución esperada bajo un modelo nulo con ausencia de competencia. Los resultados obtenidos apoyan que tanto el filtrado ambiental como la limitación a la semejanza afectan el ensamble de las comunidades vegetales de los bosques secos Tumbesinos. Un segundo tema fue identificar si la diversidad funcional está condicionada por los gradientes ambientales, y en concreto si disminuye en los ambientes más estresantes a causa del filtrado ambiental, y si por el contrario aumenta en los ambientes más benignos donde la competencia se vuelve más importante, teniendo en cuenta las posibles modificaciones a este patrón general a causa de las interacciones de facilitación. Para abordar este estudio analizamos tanto las variaciones en la diversidad funcional (respecto a los de los cinco rasgos funcionales empleados en el primer capítulo de la tesis) como las variaciones de diversidad filogenética a lo largo de un gradiente de estrés climático en los bosques tumbesinos, y se contrastaron frente a las diversidades esperadas bajo un modelo de ensamblaje completamente aleatorio de la comunidad. Los análisis mostraron que tan sólo la diversidad de tamaños foliares siguió el patrón de variación esperado, disminuyendo a medida que aumentó el estrés abiótico mientras que ni el resto de rasgos funcionales ni la diversidad funcional multivariada ni la diversidad filogenética mostraron una variación significativa a lo largo del gradiente ambiental. Un tercer tema fue evaluar si los procesos que organizan la estructura funcional de la comunidad operan a diferentes escalas espaciales. Para ello cartografié todos los árboles y arbustos de más de 5 cm de diámetro en una parcela de 9 Ha de bosque seco y caractericé funcionalmente todas las especies. Dicha parcela fue dividida en subparcelas de diferente tamaño, obteniéndose subparcelas a seis escalas espaciales distintas. Los resultados muestran agregación de estrategias funcionales semejantes a escalas pequeñas, lo que sugiere la existencia bien de filtros ambientales actuando a escala fina o bien de procesos competitivos que igualan la estrategia óptima a dichas escalas. Finalmente con la misma información de la parcela permanente de 9 Ha. Nos propusimos evaluar el efecto y comportamiento de las especies respecto a la organización de la diversidad taxonómica, funcional y filogenética. Para ello utilicé tres funciones sumario espaciales: ISAR- para el nivel taxonómico, IFDAR para el nivel funcional y IPSVAR para el nivel filogenética y las contrastamos frente a modelos nulos que describen la distribución espacial de las especies individuales. Los resultados mostraron que en todas las escalas espaciales consideradas para ISAR, IFDAR y IPSVAR, la mayoría de las especies se comportaron como neutras, es decir, que están rodeados por la riqueza de diversidad semejante a la esperada. Sin embargo, algunas especies aparecieron como acumuladoras de diversidad funcional y filogenética, lo que sugiere su implicación en procesos competitivos de limitación de la semejanza. Una pequeña proporción de las especies apareció como repelente de la diversidad funcional y filogenética, lo que sugiere su implicación en un proceso de filtrado de hábitat. En este estudio pone de relieve cómo el análisis de las dimensiones alternativas de la biodiversidad, como la diversidad funcional y filogenética, puede ayudarnos a entender la co-ocurrencia de especies en diversos ensambles de comunidad. Todos los resultados de este estudio aportan nuevas evidencias de los procesos de ensamblaje de la comunidad de los Bosques Estacionalmente secos y como las variables ambientales y la competencia juegan un papel importante en la estructuración de la comunidad. ABSTRACT The traditional concept of the rules assembly for species communities reflects the idea that species do not co-occur at random but are restricted in their co-occurrence by interspecific competition or an environmental filter. In this thesis, I addressed the importance of the se processes in the assembly of plant communities in the dry forests of southern Ecuador. This study was conducted in the biogeographic region of Tumbesina has the largest concentration of well-conserved tropical dry forests of southern Ecuador, and is recognized as one of the most important areas of endemism in the world. The climate is characterized by a dry season from May to December and a rainy season from January to April. The annual temperature varies between 20 ° C and 26 ° C and an average annual rainfall between 300 and 700 mm. I first assessed whether the distribution of functional traits at the level of the community is compatible with the existence of an environmental filter (imposed by habitat) or the existence of a limitation on functional similarity imposed by interspecific competition. This analysis was conducted for 58 species of woody plants spread over 109 plots of 10 x 50 m. Specifically, I compared the distribution of values of five functional traits (maximum height, wood density, specific leaf area, leaf size and mass of the seed), via selected statistical properties (range, variance, kurtosis and analyzed the standard deviation of the distribution of the closest functional species) distances and compared with a expected distribution under a null model of no competition. The results support that both environmental filtering and a limitation on trait similarity affect the assembly of plant communities in dry forests Tumbesina. My second chapter evaluated whether variation in functional diversity is conditioned by environmental gradients. In particular, I tested whether it decreases in the most stressful environments because of environmental filters, or if, on the contrary, functional diversity is greater in more benign environments where competition becomes more important (notwithstanding possible changes to this general pattern due to facilitation). To address this theme I analyzed changes in both the functional diversity (maximum height, wood density, specific leaf area, leaf size and mass of the seed) and the phylogenetic diversity, along a gradient of climatic stress in Tumbes forests. The observed patterns of variation were contrasted against the diversity expected under a completely random null model of community assembly. Only the diversity of leaf sizes followed the hypothesis decreasing in as trait variation abiotic stress increased, while the other functional traits multivariate functional diversity and phylogenetic diversity no showed significant variation along the environmental gradient. The third theme assess whether the processes that organize the functional structure of the community operate at different spatial scales. To do this I mapped all the trees and shrubs of more than 5 cm in diameter within a plot of 9 hectares of dry forest and functionally classified each species. The plot was divided into subplots of different sizes, obtaining subplots of six different spatial scales. I found aggregation of similar functional strategies at small scales, which may indicate the existence of environmental filters or competitive processes that correspond to the optimal strategy for these fine scales. Finally, with the same information from the permanent plot of 9 ha, I evaluated the effect and behavior of individual species on the organization of the taxonomic, functional and phylogenetic diversity. The analysis comprised three spatial summary functions: ISAR- for taxonomic level analysis, IFDAR for functional level analysis, and IPSVAR for phylogenetic level analysis, in each case the pattern of diversity was contrasted against null models that randomly reallocate describe the spatial distribution of individual species and their traits. For all spatial scales considering ISAR, IFDAR and IPSVAR, most species behaved as neutral, i.e. they are surrounded by the diversity of other traits similar to that expected under a null model. However, some species appeared as accumulator of functional and phylogenetic diversity, suggesting that they may play a role in competitive processes that limiting similarity. A small proportion of the species appeared as repellent of functional and phylogenetic diversity, suggesting their involvement in a process of habitat filtering. These analysis highlights that the analysis of alternative dimensions of biodiversity, such as functional and phylogenetic diversity, can help us understand the co-occurrence of species in the assembly of biotic communities. All results of this study provide further evidence of the processes of assembly of the community of the seasonally dry forests as environmental variables and competition play an important role in structuring the community.

10Be records of sediment cores from high northern latitudes

The 10Be records of four sediment cores forming a transect from the Norwegian Sea at 70°N (core 23059) via the Fram Strait (core 23235) to the Arctic Ocean at 86°N (cores 1533 and 1524) were measured at a high depth resolution. Although the material in all the cores was controlled by different sedimentological regimes, the 10Be records of these cores were superimposed by glacial/interglacial changes in the sedimentary environment. Core sections with high 10Be concentrations ( >1 * 10**9 at/g) are related to interglacial stages and core sections with low10Be concentrations ( <0.5 * 10**9 at/g) are related to glacial stages. Climatic transitions (e.g., Termination II, 5/6) are marked by drastic changes in the 10Be concentrations of up to one order of magnitude. The average 10Be concentrations for each climatic stage show an inverse relationship to their corresponding sedimentation rates, indicating that the 10Be records are the result of dilution with more or less terrigenous ice-rafted material. However, there are strong changes in the 10Be fluxes (e.g., Termination II) into the sediments which may also account for the observed oscillations. Most likely, both processes affected the 10Be records equally, amplifying the contrast between lower (glacials) and higher (interglacials) 10Be concentrations. The sharp contrast of high and low 10Be concentrations at climatic stage boundaries are an independent proxy for climatic and sedimentary change in the Nordic Seas and can be applied for stratigraphic dating (10Be stratigraphy) of sediment cores from the northern North Atlantic and the Arctic Ocean.