Comparing the establishment of an invasive and an endemic palm species in the Atlantic rainforest

Background: Biological invasions are one of the major causes of biodiversity loss, yet remain rather understudied in tropical environments. The Australian palm tree Archontophoenix cunninghamiana was introduced into Brazil for ornamental purposes, but has become an invasive species in urban and suburban forest patches. The substitution of A. cunninghamiana by the native palm Euterpe edulis has been proposed as a management action. Aims: We aimed to evaluate the regeneration potential of these two palm species in an Atlantic forest remnant in south-eastern Brazil where both species occur. Methods: We compared seedling establishment and seed longevity of both species through seed sowing, and also measured the contribution of A. cunninghamiana to the local seed rain and seed bank. Results: Nearly half of the non-anemochoric diaspores collected from the seed rain belonged to A. cunninghamiana, which represented a high propagule pressure in the community. The distribution of the alien palm seeds in the seed rain correlated with the distribution of nearby young and adult individuals inside the forest. Neither A. cunninghamiana nor E. edulis appeared to have a persistent seed bank in a burial experiment; seedling survival experiments suggested a much better performance for A. cunninghamiana, which had a survival rate of ca. 30% compared with a rate of only 3.5% for E. edulis. Conclusions: The results suggest a higher regeneration capacity for the alien palm over the native species when co-occurring in a forest fragment. Management actions are thus proposed to reduce a potential biological invasion process.

Molecular phylogeography of the European coastal plants Crithmum maritimum L., Halimione portulacoides (L.) Aellen, Salsola kali L. and Calystegia soldanella (L.) R. Br.

The intraspecific phylogeography of four European coastal plants, Crithmum maritimum, Halimione portulacoides, Salsola kali and Calystegia soldanella, was inferred from AFLP and ITS data. Only in C. maritimum, H. portulacoides and S. kali, a spatial genetic structure was revealed. The phylogeographic similarities and dissimilarities of these species include: (1) All three have distinct Black/Aegean and Adriatic Sea clusters. (2) Salsola kali and H. portulacoides show a distinct Atlantic/North Sea/Baltic Sea cluster, while Atlantic and eastern Spanish material of C. maritimum clustered together. (3) In the west Mediterranean, only S. kali forms a single cluster, while both H. portulacoides and C. maritimum display a phylogeographic break in the vicinity of the southern French coast. For S. kali, AFLP and ITS data concur in identifying separate Atlantic, east and west Mediterranean clades. All these patterns are postulated to result from both temperature changes during the last glacial and contemporary sea currents. No geographic AFLP structure was revealed in C. soldanella, both at the range-wide and population level. This was attributed to the remarkable seed dispersal ability of this species and possibly its longevity and clonal growth, preserving a random pattern of genetic variation generated by long-distance seed dispersal over long time periods.

European phylogeography of the coastal plants Cakile maritima Scop. (Brassicaceae) and Eryngium maritimum L. (Apiaceae)

Linear dispersal systems, such as coastal habitats, are well suited for phylogeographic studies because of their low spatial complexity compared to three dimensional habitats. Widely distributed coastal plant species additionally show azonal and often essentially continuous distributions. These properties, firstly, make it easier to reconstruct historical distributions of coastal plants and, secondly, make it more likely that present distributions contain both Quaternary refugia and recently colonized areas. Taken together this makes it easier to formulate phylogeographic hypotheses. This work investigated the phylogeography of Cakile maritima and Eryngium maritimum, two species growing in sandy habitats along the north Atlantic Ocean and the Mediterranean Sea coasts on two different spatial scales using AFLP data. The genetic structure of these species was investigated by sampling single individuals along most of their distributions from Turkey to south Sweden. On a regional scale the population genetic structure of both species was also studied in detail in the Bosporus and Dardanelles straits, the Strait of Gibraltar and along a continuous stretch of dunes in western France. Additionally, populations of C. maritima were investigated in the Baltic Sea/Kattegat/North Sea area. Over the complete sampling range the species show both differences and similarities in their genetic structure. In the Mediterranean Sea, both species contain Aegean Sea/Black Sea and west Mediterranean clusters. Cakile maritima additionally shows a clustering of Ionian Sea/Adriatic Sea collections. Further, both species show a subdivision of Atlantic Ocean/North Sea/Baltic Sea material from Mediterranean. Within the Atlantic Ocean group, C. maritima from the Baltic Sea and the most northern Atlantic localities form an additional cluster while no such substructure was found in E. maritimum. In all three instances where population genetic investigations of both species were performed in the same area, the results showed almost complete congruency of spatial genetic patterns. In the Aegean/Black Sea/Marmara region a subdivision of populations into a Black Sea, a Sea of Marmara and an Aegean Sea group is shared by both species. In addition the Sea of Marmara populations are more close to the Aegean Sea populations than they are to the Black Sea populations in both cases. Populations from the Atlantic side of the Strait of Gibraltar are differentiated from those on the Mediterranean side in both species, a pattern that confirms the results of the wide scale study. Along the dunes of West France no clear genetic structure could be detected in any of the species. Additionally, the results from the Baltic Sea/North Sea populations of C. maritima did not reveal any geographical genetic pattern. It is postulated that the many congruencies between the species are mainly due to a predominantly sea water mediated seed dispersal in both species and their shared sandy habitat. The results are compared to hypothetical distributions for the last glacial maximum based on species specific temperature requirements. It is argued that in both species the geographical borders of the clusters in the Mediterranean area were not affected by quaternary temperature changes and that the Aegean/Black Sea/Marmara cluster, and possibly the Ionian Sea/Adriatic Sea cluster in C. maritima, is the result of sea currents that isolate these basins from the rest of the sampled areas. The genetic gap in the Strait of Gibraltar between Atlantic Ocean and Mediterranean Sea populations in both species is also explained in terms of sea currents. The existence of three subgroups corresponding to the Aegean Sea, Black Sea and Sea of Marmara basins is suggested to have arisen due to geographical isolation during periods of global sea regressions in the glacials. The population genetic evidence was inconclusive regarding the Baltic Sea cluster of C. Maritima from the wide scale study. The results of this study are very similar to those of an investigation of three other coastal plant species over a similar range. This suggests that the phylo-geographic patterns of widespread coastal plants may be more predictable than those of other terrestrial plants.

Impact of forest fragmentation and disturbance on the endangered tropical tree Prunus africana (Rosaceae)

Anthropogene Fragmentierung und Störung von Wäldern beeinflussen ökologische Prozesse. Darüber hinaus werden genetische Drift und Inzucht verstärkt und die Fitness von Populationen beeinträchtigt. Um die Einflüsse von Fragmentierung und Störung auf die Biodiversität und Prozesse in tropischen Wäldern zu ermitteln, habe ich im „Kakamega Forest“, West-Kenia, die Baumart Prunus africana genauer untersucht. Dabei lag der Fokus auf (i) der Frugivorengemeinschaft und Samenausbreitung, (ii) der Kleinsäugergemeinschaft im Kontext der Samenprädation und (iii) der genetische Populationsstruktur von Keimlingen und adulten Bäumen. Der Vergleich von Keimlingen mit adulten Bäumen ermöglicht es, Veränderungen im Genfluss zwischen Generationen festzustellen. Die Ergebnisse zeigten, dass im untersuchten Waldgebiet insgesamt 49 frugivore Arten (Affen und Vögel) vorkommen. Dabei lag die Gesamtartenzahl im zusammenhängenden Wald höher als in den isoliert liegenden Fragmenten. An den Früchten von P. africana konnten insgesamt 36 Arten fressend beobachtet werden. Hier jedoch wurden in Fragmenten eine leicht erhöhte Frugivorenzahl sowie marginal signifikant erhöhte Samenausbreitungsraten nachgewiesen. Der Vergleich von stark gestörten mit weniger gestörten Flächen zeigte eine höhere Gesamtartenzahl sowie eine signifikant höhere Frugivorenzahl in P. africana in stark gestörten Flächen. Entsprechend war die Samenausbreitungsrate in stark gestörten Flächen marginal signifikant erhöht. Diese Ergebnisse deuten darauf hin, dass die quantitative Samenausbreitung in fragmentierten und gestörten Flächen etwas erhöht ist und somit eine gewisse Artenredundanz besteht, die den Verlust einzelner Arten ausgleichen könnte. Prunus africana Samen, die auf dem Boden lagen, wurden hauptsächlich von einer Nagerart (Praomys cf. jacksonii) erbeutet. Dabei war in gestörten Waldbereichen eine tendenziell höhere Prädatoraktivität zu beobachten als in weniger gestörten. Zudem waren einzelne Samen im Gegensatz zu Samengruppen in gestörten Flächen signifikant höherem Prädationsdruck ausgesetzt. Diese Ergebnisse zeigen, dass Fragmentierung sowie anthropogene Störungen auf unterschiedliche Prozesse im Lebenszyklus eines tropischen Baumes gegensätzliche Effekte haben können. Eine Extrapolation von einem auf einen anderen Prozess kann somit nicht erfolgen. Die genetische Differenzierung der adulten Baumpopulationen war gering (FST = 0.026). Der Großteil ihrer Variation (~ 97 %) lag innerhalb der Populationen, was intensiven Genfluss in der Vergangenheit widerspiegelt. Die genetische Differenzierung der Keimlinge war etwas erhöht (FST = 0.086) und ~ 91 % ihrer Variation lag innerhalb der Populationen. Im Gegensatz zu den adulten Bäumen konnte ich für Keimlinge ein „Isolation-by-distance“-Muster feststellen. Somit sind erste Hinweise auf begrenzten Genfluss im Keimlingsstadium infolge von Fragmentierung gegeben. Obwohl die Momentaufnahmen im Freiland keine Abnahme in der Frugivorenzahl und Samenausbreitung von P. africana als Folge von Fragmentierung beobachten ließen, weisen die Ergebnisse der genetischen Studie auf einen bereits reduzierten Genaustausch zwischen den Populationen hin. Somit lässt sich feststellen, dass die Faktoren Fragmentierung und Störung genetische Diversität, ökologische Prozesse und Artendiversität in Wäldern jeweils auf unterschiedliche Weise beeinflussen. Um Konsequenzen derartiger Einflüsse folgerichtig abschätzen zu können, sind Studien auf unterschiedlichen Diversitätsebenen unabdingbar.

Genetic dynamics across early life stages in the tropical tree Prunus africana (Rosaceae)

In many plant species, the genetic template of early life-stages is formed by animal-mediated pollination and seed dispersal and has profound impact on further recruitment and population dynamics. Understanding the impact of pollination and seed dispersal on genetic patterns is a central issue in plant population biology. In my thesis, I investigated (i) contemporary dispersal and gene flow distances as well as (ii) genetic diversity and spatial genetic structure (SGS) across subsequent recruitment stages in a population of the animal-pollinated and dispersed tree Prunus africana in Kakamega Forest, West Kenya. Using microsatellite markers and parentage analyses, I inferred distances of pollen dispersal (father-to-mother), seed dispersal/maternal gene flow (mother-to-offspring) as well as paternal gene flow (father-to-offspring) for four early life stages of the species (seeds and fruits, current year seedlings, seedlings ≤ 3yr, seedlings > 3yr). Distances of pollen and seed dispersal as well as paternal gene flow were significantly shorter than expected from the spatial arrangement of trees and sampling plots. They were not affected by the density of conspecific trees in the surrounding. At the propagule stage, mean pollen dispersal distances were considerably (23-fold) longer than seed dispersal distances, and paternal gene flow distances exceeded maternal gene flow by a factor of 25. Seed dispersal distances were remarkably restricted, potentially leading to a strong initial SGS. The initial genetic template created by pollination and seed dispersal was extensively altered during later recruitment stages. Potential Janzen-Connell effects led to markedly increasing distances between offspring and both parental trees in older life stages. This showed that distance and density-dependent mortality factors are not exclusively related to the mother tree, but also to the father. Across subsequent recruitment stages, the pollen to seed dispersal ratio and the paternal to maternal gene flow ratio dropped to 2.1 and 3.4, respectively, in seedlings > 3yr. The relative changes in effective pollen dispersal, seed dispersal, and paternal gene flow distances across recruitment stages elucidate the mechanisms affecting the contribution of the two processes pollen and seed dispersal to overall gene flow. Using the same six microsatellite loci, I analyzed genetic diversity and SGS across five life stages, from seed rain to adults. Levels of genetic diversity within the studied P. africana population were comparable to other Prunus species and did not vary across life stages. In congruence with the short seed dispersal distances, I found significant SGS in all life stages. SGS decreased from seed and early seedling stages to older juvenile stages, and it was higher in adults than in late juveniles of the next generation. A comparison of the data with direct assessments of contemporary gene flow patterns indicate that distance- or density-dependent mortality, potentially due to Janzen-Connell effects, led to the initial decrease in SGS. Intergeneration variation in SGS could have been driven by variation in demographic processes, the effect of overlapping generations, and local selection processes. Overall, my study showed that complex sequential processes during recruitment contribute to the spatial genetic structure of tree populations. It highlights the importance of a multistage perspective for a comprehensive understanding of the impact of animal-mediated pollen and seed dispersal on spatial population dynamics and genetic patterns of trees.

Spatial isolation and genetic differentiation in naturally fragmented plant populations of the Swiss Alps

Aims The effect Of anthropogenic landscape fragmentation on the genetic diversity and adaptive potential of plant populations is a major issue in conservation biology. However, little is known about the partitioning of genetic diversity in alpine species, which occur in naturally fragmented habitats. Here, we, investigate molecular patterns of three alpine plants (Epilobium fleischeri, Geum reptans and Campanula thyrsoides) across Switzerland and ask whether Spatial isolation has led to high levels of populations differentiation, increasing over distance, and a decrease of within-population variability. We further hypothesize that file contrasting potential for long-distance dispersal (LDD) of Seed in these Species will considerably influence and explain diversity partitioning. Methods For each study species, we Sampled 20-23 individuals from each of 20-32 populations across entire Switzerland. We applied Random Amplified Polymorphic Dimorphism markers to assess genetic diversity within (Nei's expected heterozygosity, H-e; percentage of polymorphic hands, P-P) and among (analysis of molecular variance, Phi(st)) populations and correlated population size and altitude with within-populalion diversity. Spatial patterns of genetic relatedness were investigated using Mantel tests and standardized major axis regression as well as unweighted pair group method with arithmetic mean cluster analyses and Monmonier's algorithm. To avoid known biases, We standardized the numbers of populations, individuals and markers using multiple random reductions. We modelled LDD with a high alpine wind data set using the terminal velocity and height of seed release as key parameters. Additionally, we assessed a number of important life-history traits and factors that potentially influence genetic diversity partitioning (e.g. breeding system, longevity and population size). Important findings For all three species, We found a significant isolation-by-distance relationship but only a moderately high differentiation among populations (Phi(st): 22.7, 48 and 16.8%, for E. fleischeri, G. reptans and C. thyrsoides, respectively). Within-population diversity (H-c: 0.19-0.21, P-p: 62-75%) was not reduced in comparison to known results from lowland species and even small populations with < 50 reproductive individuals contained high levels of genetic diversity. We further found no indication that a high long-distance seed dispersal potential enhances genetic connectivity among populations. Gene flow seems to have a strong stochastic component causing large dissimilarity between population pairs irrespective of the spatial distance. Our results suggest that other life-history traits, especially the breeding System, may play an important role in genetic diversity partitioning. We conclude that spatial isolation in the alpine environment has a strong influence on population relatedness but that a number of factors can considerably influence the strength of this relationship.

Tight Genetic Linkage of Prezygotic Barrier Loci Creates a Multifunctional Speciation Island in Petunia

Most flowering plants depend on animal vectors for pollination and seed dispersal. Differential pollinator preferences lead to premating isolation and thus reduced gene flow between interbreeding plant populations [1, 2, 3 and 4]. Sets of floral traits, adapted to attract specific pollinator guilds, are called pollination syndromes [5]. Shifts in pollination syndromes have occurred surprisingly frequently [6], considering that they must involve coordinated changes in multiple genes affecting multiple floral traits. Although the identification of individual genes specifying single pollination syndrome traits is in progress in many species, little is known about the genetic architecture of coadapted pollination syndrome traits and how they are embedded within the genome [7]. Here we describe the tight genetic linkage of loci specifying five major pollination syndrome traits in the genus Petunia: visible color, UV absorption, floral scent production, pistil length, and stamen length. Comparison with other Solanaceae indicates that, in P. exserta and P. axillaris, loci specifying these floral traits have specifically become clustered into a multifunctional “speciation island” [ 8 and 9]. Such an arrangement promotes linkage disequilibrium and avoids the dissolution of pollination syndromes by recombination. We suggest that tight genetic linkage provides a mechanism for rapid switches between distinct pollination syndromes in response to changes in pollinator availabilities.

Grazing response patterns indicate isolation of semi-natural European grasslands

Identifying drivers of species diversity is a major challenge in understanding and predicting the dynamics of species-rich semi-natural grasslands. In particular in temperate grasslands changes in land use and its consequences, i.e. increasing fragmentation, the on-going loss of habitat and the declining importance of regional processes such as seed dispersal by livestock, are considered key drivers of the diversity loss witnessed within the last decades. It is a largely unresolved question to what degree current temperate grassland communities already reflect a decline of regional processes such as longer distance seed dispersal. Answering this question is challenging since it requires both a mechanistic approach to community dynamics and a sufficient data basis that allows identifying general patterns. Here, we present results of a local individual- and trait-based community model that was initialized with plant functional types (PFTs) derived from an extensive empirical data set of species-rich grasslands within the `Biodiversity Exploratories' in Germany. Driving model processes included above- and belowground competition, dynamic resource allocation to shoots and roots, clonal growth, grazing, and local seed dispersal. To test for the impact of regional processes we also simulated seed input from a regional species pool. Model output, with and without regional seed input, was compared with empirical community response patterns along a grazing gradient. Simulated response patterns of changes in PFT richness, Shannon diversity, and biomass production matched observed grazing response patterns surprisingly well if only local processes were considered. Already low levels of additional regional seed input led to stronger deviations from empirical community pattern. While these findings cannot rule out that regional processes other than those considered in the modeling study potentially play a role in shaping the local grassland communities, our comparison indicates that European grasslands are largely isolated, i.e. local mechanisms explain observed community patterns to a large extent.

Structure and inferred dynamics of a large grove of Microberlinia bisulcata trees in central African rain forest: the possible role of periods of multiple disturbance events

A 272-ha grove of dominant Microberlinia bisulcata (Caesalpinioideae) adult trees greater than or equal to 50 cm stem diameter was mapped in its entirety in the southern part of Korup National Park, Cameroon. The approach used an earlier-established 82.5-ha permanent plot with a new surrounding 50-m grid of transect lines. Tree diameters were available from the plot but trees on the grid were recorded as being greater than or equal to 50 cm. The grove consisted of 1028 trees in 2000. Other species occurred within the grove. including the associated subdominants Tetraberlinia bifoliolata and T. korupensis. Microberlinia bisulcata becomes adult at a stein diameter of c. 50 cm and at an estimated age of 50 y. Three oval-shaped subgroves with dimensions c. 8 50 in x 13 50 in (90 ha) were defined. For two of them (within the plot) tree diameters were available. Subgroves differed in their scales and intensities of spatial tree patterns, and in their size frequency distributions, these suggesting differing past dynamics. The modal scale of clumping was 40-50 m. Seed dispersal by pod ejection (to c. 50 in) was evident from the semi-circles of trees at the grove's edge and from the many internal circles (100-200 m diameter). The grove has the capacity. therefore, to increase at c. 100 m per century. To form its present extent and structure. it is inferred that it expanded and infilled from a possibly smaller area of lower adult-tree density. This possibly happened in three waves of recruitment, each one determined by a period of several intense disturbances. Climate records for Africa show that 1740-50 and 1820-30 were periods of drought, and that 1870-1895 was also regionally very dry. Canopy openings allow the light-demanding and fast-growing ectomycorrhizal M. bisulcata to establish, but successive releases are thought to be required to achieve effective recruitment. Nevertheless, in the last 50 y there were no major events and recruitment in the grove was very poor. This present study leads to a new hypothesis of the role of periods of multiple extreme events being the driving factor for the population dynamics of many large African tree species such as M. bisulcata.

Genetic diversity and differentiation follow secondary succession in a multi-species study on woody plants from subtropical China

Aims: Species diversity and genetic diversity may be affected in parallel by similar environmental drivers. However, genetic diversity may also be affected independently by habitat characteristics. We aim at disentangling relationships between genetic diversity, species diversity and habitat characteristics of woody species in subtropical forest. Methods: We studied 11 dominant tree and shrub species in 27 plots in Gutianshan, China, and assessed their genetic diversity (Ar) and population differentiation (F’ST) with microsatellite markers. We tested if Ar and population specific F’ST were correlated to local species diversity and plot characteristics. Multi-model inference and model averaging were used to determine the relative importance of each predictor. Additionally we tested for isolation-by-distance and isolation-by-elevation by regressing pairwise F’ST against pairwise spatial and elevational distances. Important findings: Genetic diversity was not related to species diversity for any of the study species. Thus, our results do not support joint effects of habitat characteristics on these two levels of biodiversity. Instead, genetic diversity in two understory shrubs, Rhododendron simsii and Vaccinium carlesii, was affected by plot age with decreasing genetic diversity in successionally older plots. Population differentiation increased with plot age in Rhododendron simsii and Lithocarpus glaber. This shows that succession can reduce genetic diversity within, and increase genetic diversity between populations. Furthermore, we found four cases of isolation-by-distance and two cases of isolation-by-elevation. The former indicates inefficient pollen and seed dispersal by animals whereas the latter might be due to phenological asynchronies. These patterns indicate that succession can affect genetic diversity without parallel effects on species diversity and that gene flow in a continuous subtropical forest can be restricted even at a local scale.

Contribution of woody habitat islands to the conservation of birds and their potential ecosystem services in an extensive Colombian rangeland

The conservation of birds and their habitats is essential to maintain well-functioning ecosystems including human-dominated habitats. In simplified or homogenized landscapes, patches of natural and semi-natural habitat are essential for the survival of plant and animal populations. We compared species composition and diversity of trees and birds between gallery forests, tree islands and hedges in a Colombian savanna landscape to assess how fragmented woody plant communities affect forest bird communities and how differences in habitat characteristics influenced bird species traits and their potential ecosystem function. Bird and tree diversity was higher in forests than in tree islands and hedges. Soil depth influenced woody species distribution, and canopy cover and tree height determined bird species distribution, resulting in plant and bird communities that mainly differed between forest and non-forest habitat. Bird and tree species and traits widely co-varied. Bird species in tree islands and hedges were on average smaller, less specialized to habitat and more tolerant to disturbance than in forest, but dietary differences did not emerge. Despite being less complex and diverse than forests, hedges and tree islands significantly contribute to the conservation of forest biodiversity in the savanna matrix. Forest fragments remain essential for the conservation of forest specialists, but hedges and tree islands facilitate spillover of more tolerant forest birds and their ecological functions such as seed dispersal from forest to the savanna matrix.

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.

Estudio de la ecología de Ulmus laevis Pallas en la Península Ibérica

Esta tesis doctoral pretende profundizar en el conocimiento de la ecología de Ulmus laevis Pallas, especie autóctona en peligro de extinción en la Península Ibérica, con el fin de proponer medidas adecuadas para su conservación. Se ha estudiado la distribución natural de la especie atendiendo a aspectos edáficos. Los resultados muestran que U. laevis presenta menor capacidad de acidificación de la rizosfera, menor actividad de la reductasa férrica y menor homeostasis que U. minor Mill. cuando crecen en sustratos con una disponibilidad de hierro limitada. Estas diferencias ayudan a comprender la distribución de ambas especies en la Península Ibérica: U. laevis se ve restringido a suelos ácidos o moderadamente ácidos, mientras que U. minor es capaz de habitar tanto suelos ácidos como básicos. Se han analizado las propiedades hidráulicas y anatómicas de U. laevis, constatando que sus características son favorables en ambientes con gran disponibilidad hídrica y que se trata del olmo ibérico más vulnerable a la cavitación por estrés hídrico, por lo que la aridificación del clima y la pérdida de los freáticos supone un riesgo para sus poblaciones. Para evaluar la capacidad de recuperación de la especie se han estudiado la diversidad y estructura genética espacial de las dos mayores poblaciones españolas. Los resultados evidencian que estas poblaciones mantienen niveles de diversidad equiparables o ligeramente superiores a los europeos, pese a haber sufrido un cuello de botella prolongado durante las glaciaciones y a las reducciones poblacionales recientes. En la actualidad la endogamia no representa un riesgo para estas poblaciones. También se ha analizado la producción, dispersión y predación de semillas en Valdelatas (Madrid). Los resultados han mostrado que el viento dispersa las sámaras a corta distancia (<30 m) y que los años no veceros las probabilidades de establecimiento de regenerado son bajas. Además, la producción de sámaras vanas puede tratarse de un carácter adaptativo que aumenta la eficiencia biológica de la especie, ya que favorece la supervivencia de las semillas embrionadas disminuyendo sus tasas de predación pre- y post-dispersión. La modificación del hábitat de esta especie como consecuencia de las actividades humanas afecta de manera negativa al establecimiento del regenerado. La conservación de esta especie a largo plazo requiere la recuperación de los niveles freáticos y de regímenes hidrológicos que permitan avenidas, ya que estas crean las condiciones adecuadas para el establecimiento de regenerado al eliminar la vegetación preexistente y depositar barro. ABSTRACT Ulmus laevis Pallas is an endangered species in the Iberian Peninsula. Therefore, in order to be able to propose adequate management guidelines for its conservation, this PhD Thesis intends to advance the knowledge on the species ecology in the region. Firstly, the species natural distribution was studied in relation to soil nature. Results show that U. minor Mill. had a higher root ferric reductase activity and proton extrusion capability than U. laevis, and maintained a better nutrient homeostasis when grown under iron limiting conditions. These differences in root Fe acquisition efficiencies proved helpful to understand the distribution of these species in the Iberian Peninsula, where U. laevis is restricted to acid or moderately acid soils, whereas U. minor can grow both in acid and basic soils. Secondly, we studied Ulmus laevis’ xylem anatomy and hydraulic traits. These proved favourable for growing under high water availability, but highly susceptible to drought-stress cavitation. Therefore, this species is vulnerable to the Iberian Peninsula’s aridification. Spatial genetic structure and diversity were evaluated in two of the biggest U. laevis populations in Spain in order to evaluate their recovery capabilities. These populations maintain similar or slightly higher diversity levels than European populations, despite having undergone an ancestral genetic bottleneck and having suffered recent population size reductions. No inbreeding problems have been detected in these populations. Seed production, dispersal and predation were assessed in Valdelatas’ elm grove (Madrid). Despite U. laevis samaras being winged nuts, wind dispersed them short distances from the mother tree (<30 m). The seed shadow models show that non-mast years provide very few chances for the stand to regenerate due to their low full seed flux. Empty samaras deceive pre- and post-dispersal predators increasing full seed survival probabilities. Therefore, empty fruit production might be an adaptive trait that increases plant fitness. Finally, human-induced changes in water-table levels and river regulation may affect U. laevis seed dispersal and regeneration establishment negatively. The long-term conservation and expansion of this species in the Iberian Peninsula requires the recovery of water-tables and of natural hydrological regimes, as flooding eliminates vegetation, creating open microhabitats and deposits mud, creating the ideal conditions for seedling establishment.

Ecología de la dispersión de plantas en los bosques secos del suroccidente Ecuatoriano

La importancia del proceso de dispersión de semillas en la estructura y dinámica de los ecosistemas es ampliamente reconocida. Sin embargo, para los bosques tropicales estacionalmente secos los estudios relacionados con este proceso son aún escasos y dispersos en comparación con los bosques tropicales lluviosos. En este trabajo se estudió la importancia de los síndromes de dispersión de semillas en la estructuración de comunidades, mediante el análisis de los patrones de dispersión de semillas en el espacio y tiempo para comunidades de leñosas en los bosques secos del suroccidente Ecuatoriano. Esta área forma parte de la región Tumbesina, una de las áreas de endemismo más importantes del mundo, pero también uno de los hotspots más amenazados. El clima se caracteriza por una estación seca que va de mayo a noviembre y una estación lluviosa que se extiende desde diciembre a abril. Para toda esta zona se estima una temperatura promedio anual entre 20° y 26°C y una precipitación promedio anual entre 300 y 700 mm. El trabajo de campo se desarrolló entre febrero de 2009 y septiembre de 2012. El primer paso fue la recopilación de información sobre las especies leñosas nativas de los bosques secos del suroccidente de Ecuador, que permitiera asignar a cada especie a un síndrome de dispersión para determinar el espectro de síndromes de dispersión de semillas. Luego, utilizando la información disponible de 109 parcelas establecidas previamente a lo largo de cuatro cantones de la provincia de Loja que conservan bosques secos en buen estado, se analizó la relación entre el síndrome de dispersión y condiciones ambientales. La relación de los síndromes de dispersión con los patrones espaciales de las especies y con los patrones de la lluvia y banco de semillas se estudió dentro de una parcela permanente de 9 ha, en la Reserva Ecológica Arenillas. Dentro de esta parcela se estableció un transecto de aproximadamente 3,4 km, que se recorrió mensualmente para colectar excretas de cérvidos y analizar el rol de este grupo como dispersor de semillas. Una gran variedad de plantas en los bosques secos tropicales del suroccidente de Ecuador requirió la asistencia de animales para la dispersión de semillas. Sin embargo, un análisis del espectro de dispersión considerando no solo la riqueza, sino también la abundancia relativa de especies, permitió determinar que a pesar de la alta variedad de especies zoócoras, la mayor parte de la comunidad correspondía a individuos anemócoros, que no proveen ninguna recompensa para la dispersión por animales. Este patrón puede deberse a la abundancia relativa de hábitats adecuados para especies con diferente síndrome de dispersión. Las condiciones ambientales afectaron la estructura del espectro de dispersión en la comunidad de bosque seco neotropical estudiada. El análisis de la importancia relativa del síndrome de dispersión y de la heterogeneidad espacial en la formación de patrones espaciales de árboles adultos permitió determinar que la heterogeneidad ambiental ejercía un efecto adicional (y en algunos el único) en la formación de patrones agregados de la mayoría de especies estudiadas. Los resultados señalaron diferencias en los patrones espaciales de las especies dependiendo del síndrome de dispersión, pero también una gran variación en los patrones espaciales incluso entre especies del mismo síndrome de dispersión. El análisis simultáneo de los patrones de la lluvia de semillas y banco de semillas de una comunidad de leñosas y su relación con la vegetación establecida indicaron que la lluvia de semillas era temporalmente variable en número de especies y abundancia de semillas, y dependía del síndrome de dispersión. El síndrome de dispersión también influyó en la formación de bancos de semillas, siendo las especies con capacidad de dispersión limitada (autócoras) las de mayor riqueza de especies y abundancia de semillas. Los cérvidos también se consideraron como un elemento clave en el proceso de dispersión de semillas. Al menos ocho especies leñosas fueron dispersadas legítimamente vía endozoócora. La mayoría de las especies dispersadas presentaron diásporas sin adaptaciones obvias para la dispersión, por lo que la ingestión de semillas por cérvidos se constituye en una vía potencial para la dispersión de sus semillas a largas distancias y, con ello, mejora la posibilidad de colonizar nuevos sitios y mantener el flujo genético. Los resultados de este estudio aportan nuevas evidencias para el entendimiento de la importancia de los procesos de dispersión de semillas en la estructura de los bosques secos neotropicales. Uno de los principales hallazgos a partir de estos cuatro capítulos es que los patrones espaciales de las especies, así como las estrategias que utilizan para dispersarse y hacer frente a las condiciones adversas (es decir, lluvia o banco de semillas) llevan consigo un efecto del síndrome de dispersión, y que la intensidad ese efecto depende a la vez de las condiciones ambientales del lugar. ABSTRACT The importance of seed dispersal process in the estructuring and ecosystem dynamic is widely recongnized. However, for seasonally tropical dry forest studies related to this process are still scarce and scattered compared to tropical rain forests. The present research deals with the importance of seed dispersal syndromes as a driver in the community structure, focusing its attention to temporal and spatial patterns of seed dispersal in woody communities of seasonally dry forest at Southwestern Ecuador. This area is part of the Tumbesian region, one of the most important areas of endemism, but also one of the most threatened areas around the world. Climate is characterized by a dry season from May to November, and a rainy season from December to April. For the whole area an average temperature between 20 ° and 26 ° C, and an average annual rainfall between 300 and 700 mm are estimated. Fieldwork was carried out between February 2009 and September 2012. During a first step information about native woody species of dry forests of southwestern Ecuador was gathered, enabling to assign a dispersal syndrome to each species to determine the seed dispersal spectrum. In a second step, available information from 109 established plots along four municipalities in Loja province, which hold the highest and best conserved dry forest remanants, was analyzed to establish the relationship between dispersal syndromes and environmental conditions. The relationships between dispersal syndromes and species spatial patterns; and between dispersal syndromes and seed rain and seed bank patterns, were studied within a permanent plot of 9 ha, in the Arenillas Ecological Reserve. Within this plot one transect of approximately 3.4 km was set to collect monthly deer droppings, which were used to latter analyze the rol of this group as seed dispersers. The results showed that a large variety of plants in tropical dry forest of Southwestern Ecuador require animal assistance to dispers their seeds. However, an analysis of seed dispersal spectrum considering not only species richness, but also the relative abundance of species, allowed to determine that despite the high variety of zoochorous species, most individuals in the community corresponds to anemochoruos species. This shift may be due to the relative abundance of habitats that are suitable for species with different dispersal syndromes. Moreover, quantitative data analysis showed that environmental conditions affect the structure of seed dispersal spectrum in the studied community. The analysis of relative importance of dispersal syndrome, and the environmental heterogeneity on formation of adult trees spatial patterns, indicated that environmental heterogeneity exert an additional (or was the only) effect limiting the distribution of most species in this forest. The findings showed differences in spatial patterns related to dispersal syndrome, but also showed a large variation in spatial patterns even among species sharing the same dispersal syndrome. Simultaneous analysis of seed rain and seed bank patterns of a woody community, and their relationship with established vegetation, suggested that seed rain is temporally variable in species number and seeds abundance, and that variation is related to the dispersal syndrome. Dispersal syndrome also influenced on the formation of seed banks, being species with limited dispersal abilities (autochorous) the ones with highest species richness and seed abundance. Deer were found as a key element in the seed dispersal process. At least to eight woody species were dispersed legitimately by ingestion. Diaspores of most dispersed species had no obvious adaptations to seed dispersal, therefore, seed ingestion by deer represents a potential pathway for long-distance dispersal, and hence, improves the chances to colonizing new sites and to maintain gene flow. Overall, these results provide new evidence for understanding the importance of seed dispersal processes in the structure of Neotropical dry forests. One of the major findings from these four chapters is that spatial patterns of species, and the strategies used to disperse their seeds and to deal with the adverse conditions (i.e. seed rain or seed bank) are related with dispersal syndromes, and the intensity of that relation depends in turn, on environmental conditions.

Patrones y procesos espaciales en poblaciones y comunidades vegetales: nuevas herramientas e hipótesis

Generalmente los patrones espaciales de puntos en ecología, se definen en el espacio bi-dimensional, donde cada punto representado por el par ordenado (x,y), resume la ubicación espacial de una planta. La importancia de los patrones espaciales de plantas radica en que proceden como respuesta ante importantes procesos ecológicos asociados a la estructura de una población o comunidad. Tales procesos incluyen fenómenos como la dispersión de semillas, la competencia por recursos, la facilitación, respuesta de las plantas ante algún tipo de estrés, entre otros. En esta tesis se evalúan los factores y potenciales procesos subyacentes, que explican los patrones de distribución espacial de la biodiversidad vegetal en diferentes ecosistemas como bosque mediterráneo, bosque tropical y matorral seco tropical; haciendo uso de nuevas metodologías para comprobar hipótesis relacionadas a los procesos espaciales. En este trabajo se utilizaron dos niveles ecológicos para analizar los procesos espaciales, el nivel de población y el nivel de comunidad, con el fin de evaluar la importancia relativa de las interacciones intraespecíficas e interespecíficas. Me centré en el uso de funciones estadísticas que resumen los patrones de puntos para explorar y hacer inferencias a partir de datos espaciales, empezando con la construcción de un nuevo modelo nulo para inferir variantes del síndrome de dispersión de una planta parásita en España central. Se analizó la dependencia de los patrones espaciales tanto de los hospedantes afectados como de los no-afectados y se observó fuerte dependencia a pequeña y mediana distancia. Se utilizaron dos funciones (kernel) para simular la dispersión de la especie parásita y se identificó consistencia de estos modelos con otros síndromes de dispersión adicionalmente a la autodispersión. Un segundo tema consistió en desarrollar un método ANOVA de dos vías? para patrones de puntos replicados donde el interés se concentró en evaluar la interacción de dos factores. Este método se aplicó a un caso de estudio que consitió en analizar la influencia de la topografía y la altitud sobre el patrón espacial de un arbusto dominante en matorral seco al sur del Ecuador, cuyos datos provienen de patrones de puntos replicados basados en diseño. Partiendo de una metodología desarrollada para procesos uni-factoriales, se construyó el método para procesos bi-factoriales y así poder evaluar el efecto de interacción. Se observó que la topografía por sí sola así como la interacción con la altitud presentaron efecto significativo sobre la formación del patrón espacial. Un tercer tema fue identificar la relación entre el patrón espacial y el síndrome de dispersión de la comunidad vegetal en el bosque tropical de la Isla de Barro Colorado (BCI), Panamá. Muchos estudios se han desarrollado en este bosque tropical y algunos han analizado la relación síndrome-patrón espacial, sin embargo lo novedoso de nuestro estudio es que se evaluaron un conjunto amplio de modelos (114 modelos) basados en procesos que incorporan la limitación de la dispersión y la heterogeneidad ambiental, y evalúan el efecto único y el efecto conjunto, para posteriormente seleccionar el modelo de mejor ajuste para cada especie. Más de la mitad de las especies presentaron patrón espacial consistente con el efecto conjutno de la limitación de la dispersión y heterogeneidad ambiental y el porcentaje restante de especies reveló en forma equitativa el efecto único de la heterogeneidad ambiental y efecto único de limitación de la dispersión. Finalmente, con la misma información del bosque tropical de BCI, y para entender las relaciones que subyacen para mantener el equilibrio de la biodiversidad, se desarrolló un índice de dispersión funcional local a nivel de individuo, que permita relacionar el patrón espacial con cuatro rasgos funcionales clave de las especies. Pese a que muchos estudios realizados involucran esta comunidad con la teoría neutral, se encontró que el ensamble de la comunidad de BCI está afectado por limitaciones de similaridad y de hábitat a diferentes escalas. ABSTRACT Overall the spatial point patterns in ecology are defined in two-dimensional space, where each point denoted by the (x,y) ordered pair, summarizes the spatial location of a plant. The spatial point patterns are essential because they arise in response to important ecological processes, associated with the structure of a population or community. Such processes include phenomena as seed dispersal, competition for resources, facilitation, and plant response to some type of stress, among others. In this thesis, some factors and potential underlying processes were evaluated in order to explain the spatial distribution patterns of plant biodiversity. It was done in different ecosystems such as Mediterranean forest, tropical forest and dry scrubland. For this purpose new methodologies were used to test hypothesis related to spatial processes. Two ecological levels were used to analyze the spatial processes, at population and community levels, in order to assess the relative importance of intraspecific and interspecific interactions. I focused on the use of spatial statistical functions to summarize point patterns to explore and make inferences from spatial data, starting with the construction of a new null model to infer variations about the dispersal syndrome of a parasitic plant in central Spain. Spatial dependence between point patterns in a multivariate point process of affected and unaffected hosts were analyzed and strong dependence was observed at small and medium distance. Two kernel functions were used to simulate the dispersion of parasitic plant and consistency of these models with other syndromes was identified, in addition to ballistic dispersion. A second issue was to analyze altitude and topography effects on the spatial population structure of a dominant shrub in the dry ecosystem in southern Ecuador, whose data come from replicated point patterns design-based. Based on a methodology developed for uni-factorial process, a method for bi-factorial processes was built to assess the interaction effect. The topography alone and interacting with altitude showed significant effect on the spatial pattern of shrub. A third issue was to identify the relationship between the spatial pattern and dispersal syndromes of plant community in the tropical forest of Barro Colorado Island (BCI), Panamá. Several studies have been developed in this tropical forest and some focused on the spatial pattern-syndrome relationship; however the novelty of our study is that a large set of models (114 models) including dispersal limitation and environmental heterogeneity were evaluated, used to identify the only and joint effect to subsequently select the best fit model for each species. Slightly more than fifty percent of the species showed spatial pattern consistent with only the dispersal limitation, and the remaining percentage of species revealed the only effect of environmental heterogeneity and habitat-dispersal limitation joined effect, equitably. Finally, with the same information from the tropical forest of BCI, and to understand the relationships underlying for balance of biodiversity, an index of the local functional dispersion was developed at the individual level, to relate the spatial pattern with four key functional traits of species. Although many studies involve this community with neutral theory, the assembly of the community is affected by similarity and habitat limitations at different scales.