Modelling the spatio-temporal pattern of primary dispersal in stone pine (Pinus pinea L.) stands in the Northern Plateau (Spain)

Natural regeneration in stone pine (Pinus pinea L.) managed forests in the Spanish Northern Plateau is not achieved successfully under current silviculture practices, constituting a main concern for forest managers. We modelled spatio-temporal features of primary dispersal to test whether (a) present low stand densities constrain natural regeneration success and (b) seed release is a climate-controlled process. The present study is based on data collected from a 6 years seed trap experiment considering different regeneration felling intensities. From a spatial perspective, we attempted alternate established kernels under different data distribution assumptions to fit a spatial model able to predict P. pinea seed rain. Due to P. pinea umbrella-like crown, models were adapted to account for crown effect through correction of distances between potential seed arrival locations and seed sources. In addition, individual tree fecundity was assessed independently from existing models, improving parameter estimation stability. Seed rain simulation enabled to calculate seed dispersal indexes for diverse silvicultural regeneration treatments. The selected spatial model of best fit (Weibull, Poisson assumption) predicted a highly clumped dispersal pattern that resulted in a proportion of gaps where no seed arrival is expected (dispersal limitation) between 0.25 and 0.30 for intermediate intensity regeneration fellings and over 0.50 for intense fellings. To describe the temporal pattern, the proportion of seeds released during monthly intervals was modelled as a function of climate variables – rainfall events – through a linear model that considered temporal autocorrelation, whereas cone opening took place over a temperature threshold. Our findings suggest the application of less intensive regeneration fellings, to be carried out after years of successful seedling establishment and, seasonally, subsequent to the main rainfall period (late fall). This schedule would avoid dispersal limitation and would allow for a complete seed release. These modifications in present silviculture practices would produce a more efficient seed shadow in managed stands.

An integral model for natural regeneration of Pinus pinea L. in the Northern Plateau (Spain) = Modelo integral de regeneración natural para Pinus pinea L. en la Meseta Norte (España)

Natural regeneration is an ecological key-process that makes plant persistence possible and, consequently, it constitutes an essential element of sustainable forest management. In this respect, natural regeneration in even-aged stands of Pinus pinea L. located in the Spanish Northern Plateau has not always been successfully achieved despite over a century of pine nut-based management. As a result, natural regeneration has recently become a major concern for forest managers when we are living a moment of rationalization of investment in silviculture. The present dissertation is addressed to provide answers to forest managers on this topic through the development of an integral regeneration multistage model for P. pinea stands in the region. From this model, recommendations for natural regeneration-based silviculture can be derived under present and future climate scenarios. Also, the model structure makes it possible to detect the likely bottlenecks affecting the process. The integral model consists of five submodels corresponding to each of the subprocesses linking the stages involved in natural regeneration (seed production, seed dispersal, seed germination, seed predation and seedling survival). The outputs of the submodels represent the transitional probabilities between these stages as a function of climatic and stand variables, which in turn are representative of the ecological factors driving regeneration. At subprocess level, the findings of this dissertation should be interpreted as follows. The scheduling of the shelterwood system currently conducted over low density stands leads to situations of dispersal limitation since the initial stages of the regeneration period. Concerning predation, predator activity appears to be only limited by the occurrence of severe summer droughts and masting events, the summer resulting in a favourable period for seed survival. Out of this time interval, predators were found to almost totally deplete seed crops. Given that P. pinea dissemination occurs in summer (i.e. the safe period against predation), the likelihood of a seed to not be destroyed is conditional to germination occurrence prior to the intensification of predator activity. However, the optimal conditions for germination seldom take place, restraining emergence to few days during the fall. Thus, the window to reach the seedling stage is narrow. In addition, the seedling survival submodel predicts extremely high seedling mortality rates and therefore only some individuals from large cohorts will be able to persist. These facts, along with the strong climate-mediated masting habit exhibited by P. pinea, reveal that viii the overall probability of establishment is low. Given this background, current management –low final stand densities resulting from intense thinning and strict felling schedules– conditions the occurrence of enough favourable events to achieve natural regeneration during the current rotation time. Stochastic simulation and optimisation computed through the integral model confirm this circumstance, suggesting that more flexible and progressive regeneration fellings should be conducted. From an ecological standpoint, these results inform a reproductive strategy leading to uneven-aged stand structures, in full accordance with the medium shade-tolerant behaviour of the species. As a final remark, stochastic simulations performed under a climate-change scenario show that regeneration in the species will not be strongly hampered in the future. This resilient behaviour highlights the fundamental ecological role played by P. pinea in demanding areas where other tree species fail to persist.

A silviculture-oriented spatio-temporal model for germination in Pinus pinea L. in the Spanish Northern Plateau based on a direct seeding experiment

Natural regeneration in Pinus pinea stands commonly fails throughout the Spanish Northern Plateau under current intensive regeneration treatments. As a result, extensive direct seeding is commonly conducted to guarantee regeneration occurrence. In a period of rationalization of the resources devoted to forest management, this kind of techniques may become unaffordable. Given that the climatic and stand factors driving germination remain unknown, tools are required to understand the process and temper the use of direct seeding. In this study, the spatio-temporal pattern of germination of P. pinea was modelled with those purposes. The resulting findings will allow us to (1) determine the main ecological variables involved in germination in the species and (2) infer adequate silvicultural alternatives. The modelling approach focuses on covariates which are readily available to forest managers. A two-step nonlinear mixed model was fitted to predict germination occurrence and abundance in P. pinea under varying climatic, environmental and stand conditions, based on a germination data set covering a 5-year period. The results obtained reveal that the process is primarily driven by climate variables. Favourable conditions for germination commonly occur in fall although the optimum window is often narrow and may not occur at all in some years. At spatial level, it would appear that germination is facilitated by high stand densities, suggesting that current felling intensity should be reduced. In accordance with other studies on P. pinea dispersal, it seems that denser stands during the regeneration period will reduce the present dependence on direct seeding.

Can seed production and restricted dispersal limit recruitment in Pinus pinaster Aiton from the Spanish Northern Plateau?

Natural regeneration faces increasing difficulties in dry forests from the Mediterranean basin, including for normally well-regenerating species such as maritime pine (Pinus pinaster Aiton). In this paper, we studied female fertility, seed dispersal and spread rates in P. pinaster from the Spanish Northern Plateau, where natural regeneration failure is a main concern for forest managers. For this purpose we periodically collected data from seed traps and trees located at two core locations across several years. We found significant variation in interannual cone production, with the best seed trees being the same across years. In addition, we found highly skewed distributions of female reproductive effort and large fertility differences across stands located few kilometres away. Annual seed dispersal kernels fitted lognormal or 2Dt models depending on the stand analysed, with median dispersal distances between 14 and 25 m. Kernels fitted for maximum dispersal periods showed an outstanding intraseasonal variation of median dispersal distances, from 10 to 54 m, in association to variable patterns of rainfall and maximum wind speed. The amount of seed produced appeared to be enough to guarantee the natural regeneration of the stands during the typical 20-year regeneration period. Colonisation simulations concluded that Mediterranean maritime pine has a notable dispersion capacity, which is strongly influenced by levels of fecundity and, especially, by the number and frequency of long-distance dispersal events. The latter play a key role in tree dispersion processes through enlarging the occupied area and fostering the invasion of abandoned crop land.

Vegetation and fire history since the last glacial maximum in an inland area of the western Mediterranean Basin (Northern Iberian Plateau, NW Spain)

We reconstructed vegetation responses to climate oscillations, fire and human activities since the last glacial maximum in inland NW Iberia, where previous paleoecological research is scarce. Extremely sparse and open vegetation composed of steppic grasslands and heathlands with scattered pioneer trees suggests very cold and dry conditions during the Oldest Dryas, unsuitable for tree survival in the surroundings of the study site. Slight woodland expansion during the Bolling/Allerod was interrupted by the Younger Dryas cooling. Pinewoods dominated for most of the early Holocene, when a marked increase in fire activity occurred. Deciduous trees expanded later reaching their maximum representation during the mid-Holocene. Enhanced fire activity and the presence of coprophilous fungi around 6400-6000 cal yr BP suggest an early human occupation around the site. However, extensive deforestation only started at 4500 calyrBP, when fire was used to clear the tree canopy. Final replacement of woodlands with heathlands, grasslands and cereal crops occurred from 2700 cal yr BP onwards due to land-use intensification. Our paleoecological record can help efforts aimed at restoring the natural vegetation by indicating which communities were dominant at the onset of heavy human impact, thus promoting the recovery of currently rare oak and alder stands.

Aeolian dust in Colorado Plateau soils: Nutrient inputs and recent change in source

Aeolian dust (windblown silt and clay) is an important component in arid-land ecosystems because it may contribute to soil formation and furnish essential nutrients. Few geologic surfaces, however, have been characterized with respect to dust-accumulation history and resultant nutrient enrichment. We have developed a combination of methods to identify the presence of aeolian dust in arid regions and to evaluate the roles of this dust in ecosystem processes. Unconsolidated sandy sediment on isolated surfaces in the Canyonlands region of the Colorado Plateau differs greatly in mineralogical and chemical composition from associated bedrock, mainly aeolian sandstone. Detrital magnetite in the surficial deposits produces moderately high values of magnetic susceptibility, but magnetite is absent in nearby bedrock. A component of the surficial deposits must be aeolian to account for the abundance of magnetite, which formed originally in far-distant igneous rocks. Particle-size analysis suggests that the aeolian dust component is typically as much as 20–30%. Dust inputs have enriched the sediments in many elements, including P, Mg, Na, K, and Mo, as well as Ca, at sites where bedrock lacks calcite cement. Soil-surface biologic crusts are effective dust traps that apparently record a change in dust sources over the past several decades. Some of the recently fallen dust may result from human disturbance of land surfaces that are far from the Canyonlands, such as the Mojave Desert. Some land-use practices in the study area have the potential to deplete soil fertility by means of wind-erosion removal of aeolian silt.

Cenozoic nannofossils from the Exmouth Plateau, eastern Indian Ocean

Six sites (759-764) were drilled on the Exmouth Plateau during Ocean Drilling Program Leg 122. Nannofossilrich Cenozoic sediments were recovered at all six sites, reflecting the open-ocean conditions that prevailed over the Exmouth Plateau during the Cenozoic. Calcareous nannofossils are abundant, diverse (250 different species identified), and generally well preserved throughout the composite lower Paleocene to Quaternary section. The diversity and preservation of nannofossils permits a high degree of stratigraphic resolution at each site. Site 762 on the central part of the Exmouth Plateau contains an almost unbroken Cenozoic record (only Miocene Zones NN3, NN8, and NN10 are missing). This site may prove to be a useful Cenozoic biostratigraphic and biomagnetochronologic reference section for the eastern Indian Ocean.

Benthic foraminiferal assemblages and paleoproductivity reconstructions for the last 550 kyr of the Ontong Java Plateau, western equatorial Pacific

We analyzed foraminiferal and nannofossil assemblages and stable isotopes in samples from ODP Hole 807A on the Ontong Java Plateau in order to evaluate productivity and carbonate dissolution cycles over the last 550 kyr (kilo year) in the western equatorial Pacific. Our results indicate that productivity was generally higher in glacials than during interglacials, and gradually increased since MIS 13. Carbonate dissolution was weak in deglacial intervals, but often reached a maximum during interglacial to glacial transitions. Carbonate cycles in the western equatorial Pacific were mainly influenced by changes of deep-water properties rather than by local primary productivity. Fluctuations of the estimated thermocline depth were not related to glacial to interglacial alternations, but changed distinctly at ~280 kyr. Before that time the thermocline was relatively shallow and its depth fluctuated at a comparatively high amplitude and low frequency. After 280 kyr, the thermocline was deeper, and its fluctuations were at lower amplitude and higher frequency. These different patterns in productivity and thermocline variability suggest that thermocline dynamics probably were not a controlling factor of biological productivity in the western equatorial Pacific Ocean. In this region, upwelling, the influx of cool, nutrient-rich waters from the eastern equatorial Pacific or of fresh waters from rivers have probably never been important, and their influence on productivity has been negligible over the studied period. Variations in the inferred productivity in general are well correlated with fluctuations in the eolian flux as recorded in the northwestern Pacific, a proxy for the late Quaternary history of the central East Asian dust flux into the Pacific. Therefore, we suggest that the dust flux from the central East Asian continent may have been an important driver of productivity in the western Pacific.

Benthic foraminifera in the Tasman Sea, Emerald Basin and Challenger Plateau

Im Rahmen des TASQWA-Projektes (Quarternary Variability of Water Masses in the Southern Tasman Sea and the Southern Ocean) wurde eine erstmalige quantitative und taxonomische Bestandsaufnahme der rezenten, benthischen Tiefseeforaminiferen der Korngrößenfraktion > 250 µm in 27 Sedimentoberflächenproben aus dem austral-antarktischen Gebiet durchgeführt. Es konnten 137 Arten bestimmt werden, wobei aber keine Art dominante Anteile in den Proben erreichte. Über benthische Tiefseeforaminiferen im untersuchten Gebiet existiert kaum Literatur. Es gibt zwar aus dem 19. Jhrd. sehr gut dokumentierte Foraminiferen in diesem Bereich, diese decken aber längst nicht alle gefundenen Exemplare ab. Erst um die Jahrtausendwende beschäftigten sich Autoren wieder intensiver mit den australischen und neuseeländischen, benthischen Foraminiferen. Aber auch sie drangen nicht bis in die Tiefsee vor, sondern blieben vorwiegend im Schelfbereich. Aufgrund dieser spärlichen Literatur ist jede einzelne Art ausführlich mit Synonymieliste und Abbildung dokumentiert worden. Die PAST-Analyse generierte mit den 137 Arten und den 27 Stationen sechs Faunenvergesellschaftungen, die überwiegend bathymetrisch zoniert sind. Ab 562 m beginnt am Campbell Plateau in der Hochproduktionszone die Bulimina-Vergesellschaftung. Diese Vergesellschaftung zeichnet sich durch die höchste Individuenzahl aus. Ab 959 m findet sich die Rhizammina-Vergesellschaftung, die im Untersuchungsgebiet am weitesten verbreitet ist. Die weniger oft anzutreffende Cibicides-Vergesellschaftung läßt sich ab 1660 m Tiefe finden. Nur in einer einzigen Probe an der Tasmanschwelle in 2146 m Tiefe, tritt die Reophax-Vergesellschaftung auf, in der die Textulariina überwiegen. Die weniger oft anzutreffende Ehrenbergina-Vergesellschaftung läßt sich ab 1841 m finden. In dieser Vergesellschaftung, in der die Artenanzahl fast an das Niveau der Hochproduktionszone heranreicht, halten sich Rotaliina und Textulariina die Waage. Im Emerald Becken ab 3909 m Tiefe beginnt die Jaculella- Vergesellschaftung. Diese liegt in einem echten Hungergebiet und besteht hauptsächlich aus Textulariina. Im gesamten Untersuchungsgebiet lassen sich durch die Probenauswertung vier unterschiedliche Lebensräume (Challenger Plateau, Campbell Plateau, Emerald Becken und Tasmanschwelle) ausmachen. Da jedoch nur zwei Sedimentoberflächenproben am Challenger Plateau genommen wurden, konnte dieser Bereich nur eingeschränkt mit den anderen drei Bereichen verglichen werden. Die Foraminiferengemeinschaften des Challenger Plateaus und der Tasmanschwelle können jedoch im oberen Bereich der Wassersäule auch nur eingeschränkt miteinander verglichen werden, da man an der Tasmanschwelle Sedimentoberflächenproben erst ab 1634 m genommen hat und am Campbell Plateau Proben ab 562 m vorhanden sind. Die oberen Bereiche (ab 562 m bis ca. 1300 m) des Campbell Plateaus sind Hochproduktionsbereiche, die die höchsten Individuenzahlen pro 10 cm**3 Sediment und die höchste Artenvielfalt aufweisen. Am Südwesthang des Campbell Plateaus läßt sich eine Abfolge der verschiedenen Foraminiferenvergesellschaftungen bis hinunter in das Emerald Becken nachweisen. An der Tasmanschwelle selbst läßt sich keine ausgeprägte Hochproduktionszone erkennen. Generell gibt es hier weniger Arten und weniger Individuen pro 10 cm**3 Sediment als am Cambell Plateau. Das Emerald Becken, als tiefster Bereich des Untersuchungsgebietes und als echtes Hungergebiet, nimmt eine Sonderrolle ein.

(Table 1) Abundance of tunicate spicules along the Great Barrier Reef-Queensland Plateau transect

The abundance patterns of tunicate spicules are documented for the Pliocene-Pleistocene sediments at seven sites along the Great Barrier Reef-Queensland Plateau transect. The spatial distribution pattern indicates that tunicate spicules were limited to waters shallower than 900 m. The occurrences of tunicate spicules at Sites 822 and 823 that are deeper than 900 m are ascribed to downslope transport, and their distribution patterns can be used to monitor downslope transport processes. The first common occurrence of tunicate spicules at Sites 822 and 823 around 1.6 Ma may suggest the initiation of the central Great Barrier Reef at this time. The morphology of tunicate spicules varies greatly and appears to be gradational among different forms. Older tunicate assemblages are less diverse than those in younger sediments, presumably because of diagenesis. Tunicate spicules do not appear to be a promising biostratigraphic tool for the Pliocene-Pleistocene.