Moonlight and shelter cause differential seed selection and removal by rodents

Various environmental factors may influence the foraging behaviour of seed dispersers which could ultimately affect the seed dispersal process. We examined whether moonlight levels and the presence or absence of rodentshelter affect rodentseedremoval (rate, handling time and time of removal) and seedselection (size and species) among seven oak species. The presence or absence of safe microhabitats was found to be more important than moonlight levels in the removal of seeds. Bright moonlight caused a different temporal distribution of seedremoval throughout the night but only affected the overall removal rates in open microhabitats. Seeds were removed more rapidly in open microhabitat (regardless of the moon phase), decreasing the time allocated to seed discrimination and translocation. Only in open microhabitats did increasing levels of moonlight decrease the time allocated to selection and removal of seeds. As a result, a more precise seedselection was made under shelter, owing to lower levels of predation risk. Rodent ranking preference for species was identical between full/new moon in shelter but not in open microhabitats. For all treatments, species selection by rodents was much stronger than size selection. Nevertheless, heavy seeds, which require more energy and time to be transported, were preferentially removed under shelter, where there is no time restriction to move the seeds. Our findings reveal that seedselection is safety dependent and, therefore, microhabitats in which seeds are located (sheltered versus exposed) and moonlight levels in open areas should be taken into account in rodent food selection studies.

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.

Modelling Pinus pinea forest management to attain natural regeneration under present and future climatic scenarios

Natural regeneration-based silviculture has been increasingly regarded as a reliable option in sustainable forest management. However, successful natural regeneration is not always easy to achieve. Recently, new concerns have arisen because of changing future climate. To date, regeneration models have proved helpful in decision-making concerning natural regeneration. The implementation of such models into optimization routines is a promising approach in providing forest managers with accurate tools for forest planning. In the present study, we present a stochastic multistage regeneration model for Pinus pinea L. managed woodlands in Central Spain, where regeneration has been historically unsuccessful. The model is able to quantify recruitment under different silviculture alternatives and varying climatic scenarios, with further application to optimize management scheduling. The regeneration process in the species showed high between-year variation, with all subprocesses (seed production, dispersal, germination, predation, and seedling survival) having the potential to become bottlenecks. However, model simulations demonstrate that current intensive management is responsible for regeneration failure in the long term. Specifically, stand densities at rotation age are too low to guarantee adequate dispersal, the optimal density of seed-producing trees being around 150 stems·ha−1. In addition, rotation length needs to be extended up to 120 years to benefit from the higher seed production of older trees. Stochastic optimization confirms these results. Regeneration does not appear to worsen under climate change conditions; the species exhibiting resilience worthy of broader consideration in Mediterranean silviculture.

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.

Sobre la Arquitectura en la definición del Paisaje

Comúnmente la Arquitectura se manifiesta en los edificios como un hecho de la realidad que tiene siempre carácter contemporáneo y ése debe ser el valor supremo de un arte que no distingue entre antiguo y moderno, por afectar al presente y ser contemporáneo como experiencia. Los objetos se insertan irremediablemente en su medio e incluso llegan a definirlo en ocasiones: así, arquitectura y paisaje, aparecerán a veces confundidos con diferencias difíciles de determinar. El término “paisaje” es relativamente moderno y se deriva de ciertas representaciones gráficas o pictóricas producidas en Occidente en época algo posterior al Renacimiento. No obstante, el hecho de que una palabra se pueda escribir o se cite no quiere decir que la realidad a la que responde no pueda existir, pues lo escrito es solamente un medio de expresión, y es obvio que existen otros quizá más antiguos y de idéntica importancia, pues la propia escritura no es más que el reflejo de un fenómeno que ya se ha producido con anterioridad. Parece así que el testimonio de la realidad viene dado por distintos contextos en lo que suele llamarse “cultura” de modo que las culturas pueden tener aspectos de gran interés mediante diferentes sistemas de expresión variados sin que eso tenga que pasar forzosamente por el filtro de un alfabeto y una escritura. A tenor de los primeros descubrimientos, parece que la cuestión de escribir tuvo originalmente un carácter de constatación de apunte contable y tampoco se puede establecer con certeza si algunos utilizaron la escritura como el so-porte adecuado para referir mitos o historias, pues la Arqueología no ha proporcionado sino testimonios fragmentarios; de lo que si está repleta sin duda la historia es de apuntes contables. Otra cuestión que suscita dudas es la propia definición de escritura, pues parece que los más antiguos modos de expresión escrita se reducen a pictogramas que resultan aún indescifrables. Lenguas y toponimia son también herramientas muy útiles para bucear en el pasado, pero aún así persisten dudas en las escrituras que pro-vienen de las representaciones primordiales: la propia escritura en sus orígenes parece alimentarse de las propias intuiciones inmediatas de la representación, algo que evolucionaría representando esa realidad urgente e inmediata del control o el abastecimiento que luego se convertirían en los fragmentos de texto que han podido conservarse. Es notorio, sin embargo, que ese conjunto enorme de convenios gráficos estaba sujeto a la palabra y la desaparición de las culturas determinó también la desaparición consecuente de sus lenguas, y esos signos - fueran o no escritura - han pasado definitivamente a las zonas oscuras del conocimiento. Escribir supone también disponer de capacidad de abstracción gráfica que diferencie la palabra y la cosa, y es posible que la razón que llevara a ese descubrimiento fuera simplemente el de una economía de signos, pues escritura y lenguaje son limitados por definición, mientras que las cosas del mundo son innumerables: ningún idioma contiene todas las palabras para describir todo lo que puede aparecer. Aparentemente, ese el motivo por el cual existe un localismo – un término específico que refiere algo que existe en un lugar y no en otros - en lo que se refiere a dar el nombre a la cosa y también fuente de paradigma entre los pueblos que se consideran primitivos y los que se consideran civilizados. Debe tenerse en cuenta también que esa transposición se da en ambos sentidos, de modo que las culturas más aisladas también incorporan conceptos que carecen de una explicación racional consistente. Las mitologías son así eternas, pues sirven para explicar realidades que carecen de un entendimiento alcanzable y está también bastante claro que los lenguajes resultan ser un enigma, como realidad autónoma que queda en-cerrada y se explica en sí misma. Parece también que los primeros en escribir las consonantes aisladas fueron los pueblos semíticos occidentales de las orillas del Mediterráneo, pueblos que crearon un alfabeto silábico que llegó a ser utilizado incluso por los tartesios en el sur de la península ibérica como el primer alfabeto de toda Europa occidental. Resulta patente que el término “paisaje” se ha nutrido siempre de representaciones, bien escritas o artísticas, pero queda también claro que esas representaciones se suponen derivadas de la creencia en una idea de paisaje como algo que se ve representado en sí mismo, es decir, como la imagen de una realidad externa situada fuera del individuo. Eso es un hecho trascendente, pues el paisaje requiere lejanía de la cosa, de modo que el actor – aún sabiéndose inserto en su paisaje – es incapaz de percibirlo desde dentro. En el paisaje ocurre igual que en un teatro o en una representación: los actores son conscientes de su papel y su posible cometido, pero no son los que realmente pueden disfrutar de la eficacia o de la propia presencia de la obra, pues forman parte de ella. La idea de paisaje proviene de una lectura externa a la de los propios protagonistas del drama, de modo que si quieren ser un crítico del paisaje se debe abandonar la representación para mirar el espectáculo desde una distancia adecuada, al fin y a cabo externa. De ese modo, aparece la primera constatación del hecho del paisaje como una realidad construida por objetos y personajes, pero, sobre todo, es una realidad construida por miradas. Llama también la atención otorgada en las lecturas de los especialistas hacia esa referencia externa - artística si se quiere - sobre el término “paisaje”, de modo que la bibliografía no especializada sobre el particular siempre acaba en tratados de pintura o poesía. Parece sin embargo que el hombre y sus paisajes son realidades indisolubles desde la propia aparición de la especie, de modo que llevar la cuestión hacia términos exclusivamente esteticistas parece una posición parcial. Hombre y medio han formado siempre una sola unidad, aunque esa unidad se haya convertido en determinados casos en sinónimo de depredación y destrozo. Sin embargo, esa destrucción crea igualmente un paisaje como desolación que es resultado del propio quehacer del hombre: elementos que también poseen un contenido sustancial de memoria en los paisajes industriales como un momento de la Historia previo a la automatización y a la globalización de la vida actuales. Quizá el concepto más interesante desde el punto de vista teórico sea precisamente el de esa cualidad del paisaje como mirada, algo externo producido por el contemplador en un momento ajeno a la pertenencia, una mirada que no es tan sólo estética sino comprensiva, gozosa, activa o analítica - pues existen tantas maneras de mirar como sujetos - sin que pueda definirse con precisión esa contemplación más que en términos que alcanzan la propia individualidad. En términos poéticos, también podría establecerse como ese conjunto de miradas individuales crean también una estructura que hace que ese paisaje se valore y se comprenda, de modo que desde ese punto de vista ese paisaje supone una creación colectiva. Con respeto o como depredador, el hombre se ha instalado en su medio, y al hacerlo, ha dejado huellas dentro del propio paisaje que hacen que tome una determinada significación. Naturalmente, existe una teoría que distingue entre “país” y “paisaje”, asumiendo para el primero la noción exclusiva de territorio en la cual el hombre no tiene papel estético alguno. He intentado muchas veces comprender esa posición, sin acabar de entender el planteamiento que la sustenta: parece que la visión de la cosa estará siempre humanizada, aún en las zonas “vírgenes” o intactas, pues la propia visión hace que el objeto se modifique en su propia unidad perceptiva, creando una curiosa indeterminación que lleva a los conocidos equívocos entre imagen real y representación. Efectivamente, la visión de la cosa requiere de una cultura y unos medios que la informan, de modo que un texto, una pintura, una descripción o una fotografía quedarán ya humanizadas por el propio hecho de ser informadas, pues ello otorga una forma a priori. De ese modo, el paisaje figura inscrito en una función que establece tanto aspectos de un paisaje posible como aspectos del paisaje descrito y solamente podrá hablarse sobre la potencialidad del estado final de ese paisaje y nada del propio paisaje en sí, mientras ese paisaje no quede humanizado por el propio observador. Esta cuestión obliga a elegir una definición de paisaje alejada de presupuestos esteticistas para acercarla a los puramente culturales, de modo que no existe otra vía para la investigación que contemplar esa realidad física en función de las coordenadas del hombre que la habita, dotada del contenido correspondiente a esa revelación. Lejos de las posiciones de la geomorfología, el término “paisaje” implicará así unas condiciones determinadas de contemplación por parte de un sujeto por el cual el espectáculo queda humanizado en dicho acto.Cabe pensar también si no es cierto que todos los paisajes requieren de esa condición humanizada, pues aunque no estén habitados están siempre ocupados por esa mirada que los habita, al igual que un escenario sin público que carece de vigencia. A partir de ahí se sitúan las coordenadas en donde este trabajo configura la presencia de la arquitectura en la definición del paisaje, una premisa que parece ya venir otorgada desde el principio, pues esa misma mirada del espectador ya está dotando de un sentido de orden y jerarquía a la contemplación, unas cualidades que están en la base de toda actividad arquitectónica, De hecho la propia definición de “monumento natural” - en si misma una contradicción – expresa ese conflicto, dotado de un fenómeno de admiración y solape entre cultura y naturaleza, como conceptos enfrentados desde su origen. La conclusión sobre el dilema propuesta en la tesis no ha sido otra que suponer que esas dos realidades que son la cultura y el paisaje se han solapado desde el principio de los tiempos del hombre formando un binomio indeslindable. Se ha dicho antes que el proceso de invasión del territorio por el hombre es significativo, y esa significación es la que origina una creación autónoma que se aísla como un concepto abstracto de los entes naturales, tomándolos como material de trabajo, y estableciendo una oposición conceptual en la realidad perforada e interpretada por el hombre que viene a oponerse a lo que supone la caja cerrada e ignota del enigma del mundo. La cuestión de la significación del hombre sobre el territorio se subsume en la creación de unos productos que permanecen y que son testimonio de la propia cultura, de forma que la cantidad de rastro que el hombre deja sobre el territorio contribuye en mayor medida a la cualificación del paisaje como concepto. Eso lleva a establecer que cualquier paisaje - por el mero hecho de serlo y ser definido así – es ya cultural, puesto que está configurado por los productos de la cultura. Las palabras que puedan quedar encerradas en las piedras de los monumentos o de las ciudades son las de los hombres que trabajaron allí, y también las de los que las habitaron: más aún, el propio sentido del paisaje y su conservación vienen determinados por la presencia del hombre como único interprete de conceptos como ecología o conservación, algo que se pone de manifiesto también en la espantosa devastación que producen los fenómenos propios de la naturaleza. La historia natural, al igual que la vida, están conformadas por éxito y devastación, sin que uno y otra tengan especial preferencia, pues la preferencia se alimenta de otra clase de conceptos. La cuestión de atribuir valores morales al mundo natural es algo muy antiguo, y quizá sea la fuente o el manantial de las primeras religiones, una cuestión que se une a la indefectible noción de mortalidad que define la existencia del hombre frente a la inmanencia de la naturaleza. Esa propia naturaleza está dotada intuitivamente de un carácter “inocente” suponiendo que la inocencia es lo opuesto a la sabiduría. La cuestión es bien otra, ya que la naturaleza no posee ni siquiera algo que pueda definirse como “carácter”, en el sentido personal del término. La cuestión no cae, evidentemente, del lado de las definiciones o de las cualidades, sino del propio análisis de la realidad que el hombre va construyendo, destruyendo, embelleciendo o perjudicando para definir el territorio que habita, interponiendo a su manera recursos, instalaciones y en definitiva todos los testimonios posibles como principal manifestación de su esencia. Entre los artefactos que el hombre produce, uno de los más persistentes y más conspicuamente descritos es el de la arquitectura - entendida aquí en un sentido amplio - es decir, como el conjunto de modificaciones del espacio y del territorio. El espacio se puede modificar de muchos modos, pero en cualquiera de los casos constituye una de las huellas más características que el hombre establece como manifestación física de su propio ser discursivo. También la arquitectura ha demostrado ser una de los fenómenos más persistentes, incluso más que la propia lengua que la origina en su discurso primero antes que pueda ac-cederse a una idea sobre la conformación del material. Es paradigmático que el episodio descrito sobre la Torre de Babel en la Biblia la cuestión de la ambición de los hombres frente a Dios, representada precisamente en una arquitectura, se asimile a la discusión sobre el lenguaje primordial. La cuestión no es baladí, pues el fenómeno de la creación es algo que se concede exclusivamente a los dioses, que por esa razón habitan los territorios a los que los hombres no pueden llegar; territorios de albergue en los que las mitologías sitúan a dioses y demonios, con amplios espacios intermedios donde situar a las divinidades menores, héroes o seres híbridos en los que la coyunda de los dioses con los humanos produce sujetos que alivian el sentido de la mortalidad. El comentario del Génesis también concede un valor a la técnica, al mito de Prometeo y al fuego como creador de excelencia. Frente al progreso prometeico, se postula el valor divino, y la única forma posible de paliar ese progreso es la confusión del lenguaje, pues eso será lo que produzca la dispersión y la falta de acuerdo. La cuestión también puede explicar esa afición tan común por lo canónico en arquitectura que se mantiene a lo largo de grandes períodos, al igual que una gran máquina de inercia. Parece que los conceptos primordiales de la arquitectura basados en elementos simples como el hito, el dintel, lo recto y lo curvo, la rampa o la escalera, y el uso distinto o cualificado que se otorga a la piedra, el ladrillo, la madera o el metal como componentes primordiales del material arquitectónico se haya mantenido a lo largo de muchos milenios sin apenas cambios, al igual que ocurre con las costumbres alimenticias que siguen una progresión ascendente a través de lo crudo, lo asado y lo cocido, hasta obtener distintos grados de refina-miento, pero que siempre se sustentan en la sensación primigenia. El descubrimiento de la arquitectura proviene de un cierto manejo de las dimensiones, y consiguientemente de la geometría. Pero la geometría es cosa abstracta al igual que el lenguaje, de modo que para poder realizar arquitectura se hace necesaria esa capacidad de abstracción primera que luego permite la realización de un dispositivo. La realidad y su número exhiben un divorcio, al igual que las cosas y sus nombres exhiben el suyo, y el análisis numérico es solamente una forma de ver la realidad, una forma rigurosa y exacta – aunque parcial - que solamente representa el modelo ideal al cual la realidad se aproxima en mayor o menor medida: esa aproximación matemática hace que el universo pueda condensarse parcialmente en números, al igual que la realidad puede condensarse en nombres, pero ni el nombre ni el número reflejarán el mundo en toda su complejidad. El número es quizá solamente un medio de describir las cosas, como lo serían las formas puras que responden a una configuración matemática que pueda producirse en teoría en cualquier parte del Universo. Sin embargo, para el ejercicio de la arquitectura es preciso acudir a esa simplificación que exige la visión abstracta del plano como una sección cierta realidad como un corte abstracto del elemento considerado. Con su traza o sin ella, con la propia expresión matemática que lo describa o sin precisarla, esa intuición del plano como elemento generador del espacio es anterior a aquella expresión, al igual que el habla fue anterior a la escritura, pues solamente se produjo a través de ella y como refuerzo y sustituto de la memoria. Existen así abstracciones de la memoria que aparecen derivadas de los signos de la naturaleza aparecidos solamente de forma eventual y fragmentaría: así ocurre con la línea, el cuadrado y el círculo, formas iniciales y abstractas sonsacadas en cierta medida de la observación que dan origen a los primeros signos de la arquitectura primordial, de modo que cuadrados y círculos, elevados a prismas y superficies esféricas o esferoides aparecen en tumbas y edificios elementales de los primeros tiempos mediante una geometría primordial que se superpone al paisaje en el que se inserta. Es cierto también que esas formas se encuentran ya aproximadas en objetos que se encuentran en el medio físico, líneas en extremidades, ramas y miembros; ángulos rectos en algunos cristales que se observan mediante corte o formas circulares en astros y cráteres, pero esa realidad solamente presenta su forma aproximada y no su abstracción pura, de igual modo que el jeroglífico propondrá al ave rapaz para representar la idea de vigilancia y la buena vista, o incluso que la imagen del propio ojo sustituya la idea del Dios que todo lo ve en las culturas anti-guas. La elección fue resuelta, después de muchos intentos y aproximaciones, con la adopción del ángulo recto como un artificio fácil para el replanteo a través del triángulo sagrado 3:4:5 que ya se utilizó para construir las pirámides de Egipto, y que dio origen también a la mayor parte del urbanismo antiguo, coexistiendo con la forma circular primordial en el tipo denominado “tholo”. Aquella trama cuadrangular era uno de los patrones de relleno del espacio más comunes y compactos, y esa fue probablemente la razón por la que en tiempos muy posteriores fuera adoptada como una forma eficaz permanente de organización frente al desorden topológico que procura el conjunto de asociación de plantas circulares. Otra cuestión paradigmática es que esos conceptos primordiales e ignotos - que convergen en el mismo origen de las civilizaciones - se conviertan luego en algo canónico, a través del uso. El canon en sí mismo es algo ideal, como una norma aplicable a objetos de una realidad que ha sido creada solamente como indicio del ca-non, algo abstracto que tiene proporciones estrictas que son siempre las mismas y no obedece a criterios racionales: será absurdo sin embargo buscar el canon griego entre los templos de época como algunos lo hicieron, pues los edificios solamente se aproximan a los ejemplos canónicos y por esa razón se habla del “dórico del Partenón” que es diferente del de Egina o del de Paestum, siendo todos ellos evidentemente dóricos. Sin embargo, esa idea resulta útil al tratadista, al teórico de la arquitectura y al historiador, aun-que solamente refleje una vaga idea de lo que sucede más allá del tratado. Otra cuestión es la sutileza de los ejemplos de arquitectura, y del mismo modo que los cuerpos de los seres vivos jamás son simétricos, aunque respondan a un diseño simétrico general derivado de las condiciones de la división celular, los edificios supuestamente canónicos en los que los especialistas se inspiraron para definir los órdenes clásicos no disponen de esa simetría modular exacta, sino que adaptan un modelo general al lugar siempre cambiante del emplazamiento, es decir, se adaptan con habilidad a la vez que configuran el paisaje en el que se insertan. La luz de los distintos intercolumnios del Partenón es ligeramente distinta, aunque guarde un evidente sentido de simetría axial, de manera que aquellos “órdenes” que formaron la Teoría de la Arquitectura no son más que una bella interpretación sectorial de la arquitectura antigua elaborada por los tratadistas del Renacimiento y, posteriormente, por los neoclásicos. Parece, sin embargo, que ese ansia por el canon como claridad de lenguaje es algo consustancial al desarrollo de la arquitectura, una lingua franca que tiende a evitar la dispersión producida entre los mortales por los dioses antiguos, de modo que si no era posible llegar al cielo de los dioses se procuró que el lenguaje de la Tierra fuera al menos inteligible para poder entenderse entre los hombres. Parece que la estructura del poder siempre requirió de un determinado nivel de organización, y también que las instrucciones se entendieran con claridad: parece que en nuestros tiempos esos antiguos cánones se han sustituido por la obediencia a normas abstractas, dictadas por instituciones también algo abstractas y que tienen nombres divertidos compuestos por siglas, aunque no se conozca bien su virtud. El canon actual está constituido por “normas” que dejan tranquilos a algunos, aunque parece quizá que todo ese entramado formal que sirve como anestesia para el cuerpo social procura también la destrucción de los bosques en formas de montañas ingentes de papel burocrático. Durante muchos siglos las normas fueron otras, en la forma de un canon del cual nadie podía escapar: aún así, mediante la utilización de cánones o sin ellos, la arquitectura prosperó en la civilización desde los primeros refugios cavernarios o los abrigos primigenios y de ese modo fue configurando la realidad, es decir, el paisaje. Como antes se dijo, ese es un viaje de ida y vuelta en el cual ambos se confundían y subsumían: el manejo de las formas y lenguajes de la arquitectura posibilitaría con el tiempo la distinción entre el campo en donde reina el caos y la ciudad, en donde reina teóricamente el orden, creando un divorcio que duraría milenios y que aún persiste. Esa oposición generaría también una forma de paisaje y una serie de usos simbólicos y sagrados de los elementos de la arquitectura - como son puertas y murallas - que se han mantenido conceptualmente aunque hoy las ciudades ya no posean murallas físicas, ni puertas que se cierran con la llegada de la noche. En ese sentido, la arquitectura ha podido definir el paisaje, entendiendo la arquitectura no solamente como los edificios en sí, sino como el hecho de la manifestación del hombre sobre el territorio, de modo que no podrá hablarse de paisaje sin su arquitectura, del mismo modo que no puede hablarse de paisaje sin hombres. Por esta razón, la Tesis habla sobre Arquitectura y Paisaje, pero más particularmente sobre el hecho de la arquitectura en la definición del paisaje, es decir, de como los hechos arquitectónicos van a determinar o no la cualidad y la calificación de un paisaje. Deberá partirse en primer lugar de una definición de lo que se entiende y se ha entendido comúnmente por paisaje: igualmente, y habida cuenta del contexto en el que sitúa el propio trabajo de tesis, la cuestión solamente se circunscribirá a lo que se entiende como cultura occidental, pues el desarrollo de la civilización presenta siempre un color local que hace que el análisis de un fenómeno de esta envergadura tenga distintas connotaciones en las distintas áreas de civilización. De igual modo, y habida cuenta también que el paisaje se construye a través de todas las manifestaciones culturales, se hará a veces necesario indagar en otras disciplinas no arquitectónicas para comprender el alcance de algunos casos en los cuales los restos arquitectónicos han desaparecido, o en los que subsisten escasas trazas. Una definición tan amplia de la Arquitectura llevaría a escribir un relato sobre toda la cultura occidental y por ese motivo solamente se han esbozado ideas sobre la aparición de esos signos sobre el paisaje basados en elementos antiguos y primigenios que se repiten con insistencia y van dando lugar al variado repertorio de la arquitectura occidental cómo conformación de ideas sobre el me-dio y sobre el mundo que se percibe y se habita. ABSTRACT About Architecture in defining Landscape. Abstract Architecture is commonly manifested in buildings as a fact of reality that has always a contemporary character and that should be the highest value of an art that does not distinguish between ancient and modern, to affect the present and be contemporary as experience. Objects are inserted irremediably in their midst and even sometimes come to define it: thus, architecture and landscape, appear sometimes confused with differences difficult to determine. However, the term "landscape" is relatively modern and is derived from certain graphical or pictorial representations produced in the West in some subsequent period to the Renaissance. The fact that a word can be written or quoting does not mean that the reality that can not be answered, because the writing is only a mean of expression, and it is obvious that there are other and perhaps older equally important, because the writing itself is nothing more than the reflection of a phenomenon that has occurred before. It thus appears that the testimony of reality is given by different contexts in what is called "culture", so that cultures can have aspects of great interest by many different expression systems without this necessarily have to pass through the filter of alphabet and writing. Under the initial findings, it appears that the question of writing originally had a character and finding accounting entries, and it can not be established with certainty whether some used writing as the support appropriate to refer myths or stories, because archaeology has provided only fragmentary evidence. Another issue that raises questions is what can properly be defined as writing, it seems that the oldest modes are reduced to writing pictograms are still indecipherable. Languages and place names are also very useful tools for diving in the past, but still questions remain in the scriptures that come from the primordial representations: either it is very well defined what the own writing in its origins: the beginnings seem to feed on immediate intuitions of representation, which would evolve representing reality that urgent and immediate control or supply which is then inherited into fragments. It is noteworthy, however, that this huge set of graphics agreements was subject to the word and the disappearance of cultures determined also the consequent disappearance of their languages, and those signs - whether or not they write - have passed definitively to dark areas of knowledge. Writings supposed also the capacity of abstraction graph differentiates the word and the thing, and it is possible that the reason to carry this discovery was simply that of an economy of signs, for writing and language are limited by definition, while the things of the world are innumerable: no language contains all words to describe all that may appear. Apparently, that's why there is a localism - a specific term that refers to something that exists in one place and not in others - in regards to name the thing and also the source of paradigm among peoples are considered primitive and civilized. It should be noted also that transposition occurs in both directions, so that the most isolated cultures also incorporate concepts that lack a consistent rational explanation. Mythologies are eternal and therefore serve to explain realities that lack an understanding achievable and also pretty clear that languages happen to be an enigma, as an autonomous reality that is enclosed and explains itself. It also seems that the first to write consonants were isolated western Semitic peoples from the shores of the Mediterranean, peoples who created a syllabic alphabet came to be used even by tartesios in southern Iberia as the first alphabet in Western Europe. It is clear that the term "landscape" has always nurtured representations, either written or artis-tic, but it is also clear that these representations are assumed arising from belief in an idea of landscape as something that is represented in itself, as the image of a reality external located outside the individual. That is an important fact because the landscape requires remoteness of the thing, so that the actor - even knowing insert in landscape - is unable to perceive from within. The landscape is just as in a theatre or a performance: the actors are aware of their role and their possible role, but they are not the ones who can really enjoy the efficiency or the presence of the work itself, as part of it. The idea comes from an external landscape reading the principles of players in the drama, so if you want to be a critic of the landscape should leave the actual representation to watch the spectacle from a safe distance, finally external. Thus, the first finding of fact of the landscape appears as a reality constructed by objects and characters, but above all, a reality constructed by looks. Also noteworthy given the readings of specialists to the external reference - art if it could be - on the term "landscape", so no specialized literature on the subject always ends in treatises on painting or poetry. It seems however that the man and his landscapes are inseparable realities from the very onset of the species, so bring the issue into terms exclusively aesthetics seems a partial position. Man and environment have always been a single unit, but that unit has become synonymous with certain cases predation and destruction. Nevertheless, this destruction also creates a landscape as desolation that results from proper task of man elements that also have substantial contents of memory in industrial landscapes as a time of pre-automation history and globalization of current life. Perhaps the most interesting from a theoretical point of view is precisely that quality of landscape as something external produced by the viewer in a strange time of membership, a look that is not only aesthetic but sympathetic, joyful, active concept or analytical - because there are so many ways to look as subjects - it may not be precisely defined that contemplation rather than in terms that reach one's individuality. In poetic terms, it could also be set as the set of individual gazes also create a structure that makes this landscape is valued and understood, so from that point of view that landscape is a collective creation. With respect or as a predator, man has settled in his environment and in doing so has left traces in the landscape itself that make take a certain significance. Naturally, there is a theory that distinguishes what is "home" and what is "nature" providing for the first notion of the exclusive territory in which man has no aesthetic role. I tried many times to understand this position, without understanding the approach that supports: it seems that the vision of the thing is always humane, even in the "virgin" or untouched areas, as the vision itself makes the object modified in its own perceptual unit, creating a curious indeterminacy leading to the known misunderstandings between real image and representation. Indeed, the vision of the thing requires a culture and means that the report, so that a text, a picture, a description or photograph will be humanized by the very fact of being informed, as this provides a way a priori. Thus, the landscape provides a function that sets both aspects of a potential landscape as described aspects of the landscape and can only talk about the potential of the final state of the landscape, while the landscape remains humanized by the observer himself. This question forces to choose a definition of remote landscape budgets purely cultural, so there is another way for research to contemplate that physical reality in terms of the coordinates of the man who inhabits gifted content corresponding to that revelation. Far from the positions of the geomorphology, the term "landscape" and involve a certain condition by contemplation of a subject for which the show is humanized in the act. It should also consider, in the light of the above, if it is not true that all landscapes require that humanized condition, because although they are not inhabited they are always occupied by the look that dwells, like a stage without audience produces no consistent effect. From there the coordinates where this work sets the presence of architecture in defining landscape, a premise which seems to come from the beginning given lie, because that same look is already giving the viewer a sense of order and hierarchy to contemplation, qualities that are at the basis of all architectural activity, in fact the very definition of "natural monument" - in itself a contradiction - expresses this conflict, which has a phenomenon of admiration and overlap between culture and nature as concepts faced since its inception. The conclusion on the dilemma proposed in the thesis has not been another to assume that these two realities are the culture and landscape have overlapped since the beginning of man time forming a pair. It has been said before that the process of invasion of the territory by man is significant, and that meaning is the originating autonomous creation that is isolated as an abstract concept of nature, taking them as working material, and establishing a conceptual opposition in reality and punched by the man who comes to oppose representing the closed and unknown to the enigma of the world interpreted. The question of the significance of the man on the land is subsumed in the creation of products that remain and are testimony of their own culture, so that the trace amount that the man leaves the territory contributes most to the qualification landscape as a concept. That brought into any landscape - by the mere fact of being and being well defined - is already cultural, as it is configured by culture products. The words that can be locked in the stones of the monuments or cities are those of the men who worked there, and also of those who inhabited: even more, the sense of the landscape itself and its conservation are determined by the presence of man as the sole interpreter of concepts such as ecology or conservation, something which becomes manifest also in the awful devastation that produce the phenomena of nature. The natural history, like life, are shaped by success and devastation without special preference, the preference is used for feeding on other kinds of concepts. The question of moral values attributed to the natural world is very ancient, and may be the source or the source of the first religions, an issue that joins the unfailing notion of mortality that defines the existence of man against immanence of nature. That nature is endowed intuitively an "innocent" character assuming that innocence is the opposite of wisdom. The other issue is well, since nature does not even have what is defined as "character" because that is something that serves to qualify beings, but not objects. The question does not fall clearly on the side of the definitions or qualities, but from the analysis of the reality that man is building, destroying or damaging to define the territory, interposing his way resources facilities and possible witness as the main manifestation of its essence. Among the artifacts that man produces one of the most persistent and most conspicuously disclosed is architecture as a way of modification of space and territory. The space can be modified in many ways, but in either case is one of the most characteristic traces that man establishes as a physical manifestation of his own discourse being. Architecture has also proven to be one of the most persistent phenomena, even more than their own language that originates in his speech first. The paradigm wrote in the episode described on the Tower of Babel in the Bible shows the question of ambition of men before God - represented precisely in architecture - is assimilated to the discussion about the primary language. The issue is not trivial, since the phenomenon of creation is something that is granted exclusively to the gods, for that reason inhabit the territories to which men can not reach; territories where the hostel located mythologies gods and demons, with large gaps where to place the minor deities, heroes or hybrid beings where the yoke of the gods with human subjects produces relieving sense of mortality. The commentary on Genesis also gives a value to the art, the myth of Prometheus and fire as creator of excellence. In front of promethean progress, divine value is postulated, and the only possible way to alleviate this progress is the confusion of language, because that is what will produce the dispersion and lack of agreement. The issue may also explain such a common passion for the canonical architecture maintained throughout long periods, like a great machine inertia. It seems that the main concepts of architecture based on simple elements like milestone, lintels, straight and curved stairs or ramps, or other qualified and used are granted to the stone, brick, wood or metal as the primary components of the architectural material maintained throughout many millennia are almost unchanged, as is the case with the eating habits that follow a progression through the raw, the cooked and roasted or fried until obtain different degrees of refinement, but always are based on the primal feeling. The discovery of the architecture comes from a management dimensions, and consequently the geometry. But the geometry is abstract thing like the language so that to make architecture that first absorption capacity which then allows the realization of a device is necessary. Reality and its number exhibit a divorce, like things and their names displayed his; numerical analysis is just one way of seeing reality, rigorous and accurate - though partial - only represents the ideal model to which reality is coming to a greater or lesser extent: the mathematical approach makes the universe can condense on numbers, like reality may condense on names, but neither the name nor the number will reflect in all its complexity. The number is only a means of describing things, such as the pure forms that match setup a mathematical theory that occurs anywhere in the universe. However, for the practice of architecture is necessary to go to the simplification that requires abstract view of a section plane of reality, as an abstract element considered cutting. With its trace or not, with the mathematical expression that describes what or without clarify, this intuition of the plane as a generator of space predates his own expression, like speech preceded the writing, which only occurred through it and as reinforcement and replacement of memory. There are abstractions of memory displayed on the signs of nature only on casual and fragmentary, such as line, square and circle: those initials and abstract forms located in abstraction give rise to the first signs of primordial architecture, so that squares and circles, lifting two prisms and spheroids or spherical surfaces appear in tombs and elementary buildings the first few times, and that primordial geometry overlaps the landscape in which it is inserted. It is also true that these forms are approximate objects found in the physical environment, limb lines, branches and limbs; straight in some crystals angles observed by cutting or circular in stars and craters forms, but only the approximate shape and not its abstraction, just as the hieroglyphic of a falcon to represent the idea of surveillance presents the good view, or even the image of the eye itself replace the idea of the all-seeing God. The election was resolved, after many attempts and approaches, with the adoption of the right angle as an easy trick for stake through the sacred triangle 3:4:5 already used to build the pyramids of Egypt, and also gave rise to most of the old urbanism tend coexist with the primary circular form type called "Tholo". That frame homer was one of the fill patterns of common and compact space, and that was probably the reason why in much later times was adopted as a permanent effective form of organization against the topological disorder that seeks the set of association circular plants. Another issue is that these paradigmatic and unknown primary concepts - that converge at the origin of civilizations - then become something canon, through use. The canon itself is something ideal, as a rule for objects of a reality that has been created only as an indication of the ca-non, something abstract that has strict proportions are always the same and not due to rational criteria: be absurd however seek the Greek canon among the temples of time as some did, because the buildings only approximate the canonical examples and for that reason we speak of "doric from Parthenon" which is different from Aegina or Paestum, being all clearly doric. However, this idea is useful to scholar, the architectural theorist and historian, although it reflects only a vague idea of what happens beyond the book. Another issue is the subtlety of the examples of architecture, just as the bodies of living beings are never symmetrical, even if they involve a general symmetrical design derived from the conditions of cell division, the supposedly canonical buildings where specialists were inspired to define the classical orders do not have that exact modular symmetry, but a general model adapted to the ever changing location of the site shaping the landscape in which they are inserted. The light of the various bays of the Parthenon is slightly different, but keep a clear sense of axial symmetry, so that those "orders" that formed the theory of architecture are just a beautiful sectoral interpretation of ancient architecture developed by writers of the Renaissance and later by neoclassical. It seems, however, that craving for clarity of language canon as is inherent to the development of architecture, a lingua franca that tends to avoid scattering among mortals by the ancient gods, so if it was not possible the heaven of the gods sought the language of the Earth was at least intelligible to be understood. Power structure has always required a certain level of organization, and the instructions are clearly understood: it seems that in our times these ancient canons were replaced by obedience to abstract rules, issued by institutions also somewhat abstract and have funny names made up acronym, although not well known virtue. The current canon consists of "rules" that leave some quiet, although it seems that maybe all that interweaves-do formally serving as anaesthesia for the social body also seeks the destruction of forests in forms of huge mountains of bureaucratic paper. For many centuries were other rules, in the form of a canon which no one could escape: still using royalties or without them, civilization flourished in architecture from the earliest cave shelters or shelters and the primordial reality was setting mode in landscape. As noted above, this is a return trip in which both confused and subsumed: the management of forms and languages of architecture over time would allow the distinction between the field where chaos reigns and city, where order reigns theoretically creating a divorce that lasted millennia and still persists. This opposition also generate a form of landscape and a series of symbolic and sacred uses of architectural elements - such as doors and walls - which have remained conceptually although today the cities no longer having physical walls or doors close during the night. In this sense, the architecture could define the landscape, architecture is not only understood as the buildings themselves, but the fact of the manifestation of the man on the premises, so you can not speak without his landscape architecture, the same so we can not speak of landscape without men. For this reason, the thesis discusses architecture and landscape, but more particularly to the fact of architecture in defining landscape, as the facts of architectural or not will determine the quality and qualification of a landscape. One should begin first a definition of what is understood and has been commonly understood by landscape: equally, and given the context in which it places the own thesis work, the issue only be limited to what is understood as western culture, for the development of civilization always has a local colour that makes the analysis of a phenomenon of this magnitude have different connotations in different areas of civilization. Similarly, and given also that the landscape is constructed through all cultural manifestations, will sometimes necessary to investigate other non-architectural disciplines to understand the scope of some cases in which the architectural remains have disappeared, or the remaining few traces. Such a broad definition of architecture take to write a story about all of Western culture and for this reason only been sketched ideas about the appearance of these signs on the landscape based on ancient and primitive elements are repeated insistently and leading the varied repertoire of Western architecture shaping ideas about how the media and the world is perceived and inhabited.

Ulmus laevis in the Iberian Peninsula: a review of its ecology and conservation

European white elm (Ulmus laevis Pallas) populations are scarce, small and fragmented in the Iberian Peninsula. Due to these characteristics the indigenous status of the species in the region has been questioned, whilst the species? role in Iberian riparian forest ecology has been neglected. Herein we review past studies regarding this species? distribution and ecology in the Iberian Peninsula, with special emphasis on the establishment of conservation priorities. We first present a collection of palaeogeographic, historic and genetic data suggesting that the Iberian Peninsula was a glacial refuge for U. laevis. Secondly, we analyse U. laevis distribution in relation to soil physico- chemical properties and water availability in Spain. Following this, we focus on the reproductive biology of the species, and investigate the effect of masting and empty seed production on predation and regeneration establishment. Finally, based on this knowledge, we propose conservation policies for U. laevis in the Iberian Peninsula.

What initiates speciation in passion-vine butterflies?

Studies of the continuum between geographic races and species provide the clearest insights into the causes of speciation. Here we report on mate choice and hybrid viability experiments in a pair of warningly colored butterflies, Heliconius erato and Heliconius himera, that maintain their genetic integrity in the face of hybridization. Hybrid sterility and inviability have been unimportant in the early stages of speciation of these two Heliconius. We find no evidence of reduced fecundity, egg hatch, or larval survival nor increases in developmental time in three generations of hybrid crosses. Instead, speciation in this pair appears to have been catalyzed by the association of strong mating preferences with divergence in warning coloration and ecology. In mate choice experiments, matings between the two species are a tenth as likely as matings within species. F1 hybrids of both sexes mate frequently with both pure forms. However, male F1 progeny from crosses between H. himera mothers and H. erato fathers have somewhat reduced mating success. The strong barrier to gene flow provided by divergence in mate preference is probably enhanced by frequency-dependent predation against hybrids similar to the type known to occur across interracial hybrid zones of H. erato. In addition, the transition between this pair falls at the boundary between wet and dry forest, and rare hybrids may also be selected against because they are poorly adapted to either biotope. These results add to a growing body of evidence that challenge the importance of genomic incompatibilities in the earliest stages of speciation.

Estimating the relative roles of top-down and bottom-up forces on insect herbivore populations: A classic study revisited

Although most ecologists agree that both top-down and bottom-up forces (predation and resource limitation, respectively) act in concert to influence populations of herbivores, it has proven difficult to estimate the relative contributions of such forces in terrestrial systems. Using a combination of time–series analysis of population counts recorded over 16 years and experimental data, we present the first estimates of the relative roles of top-down and bottom-up forces on the population dynamics of two terrestrial insect herbivores on the English oak (Quercus robur). Data suggest that temporal variation in winter moth, Operophtera brumata, density is dominated by time-lagged effects of pupal predators. By comparison, spatial variation in O. brumata density is dominated by host–plant quality. Overall, top-down forces explain 34.2% of population variance, bottom-up forces explain 17.2% of population variance, and 48.6% remains unexplained. In contrast, populations of the green oak tortrix, Tortrix viridana, appear dominated by bottom-up forces. Resource limitation, expressed as intraspecific competition among larvae for oak leaves, explains 29.4% of population variance. Host quality effects explain an additional 5.7% of population variance. We detected no major top-down effects on T. viridana populations. An unknown factor causing a linear decline in T. viridana populations over the 16-year study period accounts for most of the remaining unexplained variance. We discuss the observed differences between the insect species and the utility of time–series analysis as a tool in assessing the relative importance of top-down and bottom-up forces on herbivore populations.

Harvested populations are more variable only in more variable environments

Acknowledgments This work was funded by NERC grant NE/C510467/1 (T. G. Benton and S. B. Piertney) and a University of Leeds Faculty Postdoctoral Fellowship (T. C. Cameron). Data Accessibility The original time series and body size data from these experiments are available to download from DRYAD entry number http://dx.doi.org/10.5061/dryad.bq135.

A test of alternative models of diversification in tropical rainforests: Ecological gradients vs. rainforest refugia

Comparison of mitochondrial and morphological divergence in eight populations of a widespread leaf-litter skink is used to determine the relative importance of geographic isolation and natural selection in generating phenotypic diversity in the Wet Tropics Rainforest region of Australia. The populations occur in two geographically isolated regions, and within each region, in two different habitats (closed rainforest and tall open forest) that span a well characterized ecological gradient. Morphological differences among ancient geographic isolates (separated for several million years, judging by their mitochondrial DNA sequence divergence) were slight, but morphological and life history differences among habitats were large and occurred despite moderate to high levels of mitochondrial gene flow. A field experiment identified avian predation as one potential agent of natural selection. These results indicate that natural selection operating across ecological gradients can be more important than geographic isolation in similar habitats in generating phenotypic diversity. In addition, our results indicate that selection is sufficiently strong to overcome the homogenizing effects of gene flow, a necessary first step toward speciation in continuously distributed populations. Because ecological gradients may be a source of evolutionary novelty, and perhaps new species, their conservation warrants greater attention. This is particularly true in tropical regions, where most reserves do not include ecological gradients and transitional habitats.

Paleoindian large mammal hunters on the plains of North America

From ≈11,200 to 8,000 years ago, the Great Plains of North America were populated by small Paleoindian hunting groups with well developed weaponry and the expertise to successfully hunt large mammals, especially mammoths and bison. Mammoths became extinct on the Plains by 11,000 years ago, and, although paleoecological conditions were worsening, their demise may have been hastened by human predation. After this, the main target of the Plains Paleoindian hunters consisted of subspecies of bison, Bison antiquus and Bison occidentalis. As bison populations gradually diminished, apparently because of worsening ecological conditions, by ≈8,000 years ago, human subsistence was forced into a greater dependence on small animal and plant foods. Human paleoecology studies of the Paleoindian time period rely heavily on multidisciplinary efforts. Geomorphologists, botanists, soil scientists, palynologists, biologists, and other specialists aid archaeologists in data recovery and analysis, although, with few exceptions, their contributions are derived from the fringes rather than the mainstream of their disciplines.

Female choice increases offspring fitness in an arctiid moth (Utetheisa ornatrix)

In Utetheisa ornatrix (Lepidoptera, Arctiidae), the female mates preferentially with larger males. Having a larger father results in the eggs being more richly endowed with defensive pyrrolizidine alkaloid (which the female receives from the male with the sperm package, in quantity proportional to the male's body mass, and passes on to the eggs); having a larger father also results in the sons and daughters themselves being larger (body mass is heritable in Utetheisa). We provide evidence herein that these consequences enhance the fitness of the offspring. Eggs sired by larger males are less vulnerable to predation (presumably because of their higher alkaloid content), whereas sons and daughters, by virtue of being larger, are, respectively, more successful in courtship and more fecund. The female Utetheisa, therefore, by being choosy, reaps both direct phenotypic and indirect genetic benefits.

Benthic–pelagic links and rocky intertidal communities: Bottom-up effects on top-down control?

Insight into the dependence of benthic communities on biological and physical processes in nearshore pelagic environments, long considered a “black box,” has eluded ecologists. In rocky intertidal communities at Oregon coastal sites 80 km apart, differences in abundance of sessile invertebrates, herbivores, carnivores, and macrophytes in the low zone were not readily explained by local scale differences in hydrodynamic or physical conditions (wave forces, surge flow, or air temperature during low tide). Field experiments employing predator and herbivore manipulations and prey transplants suggested top-down (predation, grazing) processes varied positively with bottom-up processes (growth of filter-feeders, prey recruitment), but the basis for these differences was unknown. Shore-based sampling revealed that between-site differences were associated with nearshore oceanographic conditions, including phytoplankton concentration and productivity, particulates, and water temperature during upwelling. Further, samples taken at 19 sites along 380 km of coastline suggested that the differences documented between two sites reflect broader scale gradients of phytoplankton concentration. Among several alternative explanations, a coastal hydrodynamics hypothesis, reflecting mesoscale (tens to hundreds of kilometers) variation in the interaction between offshore currents and winds and continental shelf bathymetry, was inferred to be the primary underlying cause. Satellite imagery and offshore chlorophyll-a samples are consistent with the postulated mechanism. Our results suggest that benthic community dynamics can be coupled to pelagic ecosystems by both trophic and transport linkages.

Directed seed dispersal by bellbirds in a tropical cloud forest

A fundamental goal of plant population ecology is to understand the consequences for plant fitness of seed dispersal by animals. Theories of seed dispersal and tropical forest regeneration suggest that the advantages of seed dispersal for most plants are escape from seed predation near the parent tree and colonization of vacant sites, the locations of which are unpredictable in space and time. Some plants may gain in fitness as a fortuitous consequence of disperser behavior if certain species of dispersers nonrandomly place seeds in sites predictably favorable for seedling establishment. Such patterns of directed dispersal by vertebrates long have been suggested but never demonstrated for tropical forest trees. Here we report the pattern of seed distribution and 1-year seedling survival generated by five species of birds for a neotropical, shade-tolerant tree. Four of the species dispersed seeds to sites near the parent trees with microhabitat characteristics similar to those at random locations, whereas the fifth species, a bellbird, predictably dispersed seeds under song perches in canopy gaps. The pattern of seedling recruitment was bimodal, with a peak near parent trees and a second peak, corresponding to bellbird song perches, far (>40 m) from parent trees. Seedling survival was higher for seeds dispersed by bellbirds than by the other species, because of a reduction in seedling mortality by fungal pathogens in gaps. Thus, bellbirds play a significant role in seed dispersal by providing directed dispersal to favorable sites and therefore may influence plant recruitment patterns and species diversity in Neotropical forests.

Multiple benefits of multiple mating in guppies

The rewards of promiscuity for males are undisputed. But why should a female mate promiscuously, particularly when her partners offer no resources other than sperm and increase her chances of succumbing to predation or disease? This question has been hotly debated but at present remains largely unresolved [Jennions, M. D. & Petrie, M. (2000) Biol. Rev. 75, 21–64]. One possibility is that females exploit postcopulatory mechanisms, such as sperm competition, to increase both the quality and quantity of their offspring. In this paper, we use the Trinidadian guppy, a species with a resource-free mating system, to test the hypothesis that females gain multiple benefits from multiple mating. Our results indicate that multiply mated females secure substantive advantages: They have shorter gestation times and larger broods, and they produce offspring with better developed schooling abilities and escape responses than their singly mated counterparts.