Caracterización morfofisiológica de semillas de especies leñosas distribuidas en dos zonas secas presentes en el Sur del Ecuador

Sudamérica es una de las zonas con mayor cantidad de bosque seco tropical a nivel mundial. No obstante, estos bosques han sido poco conocidos y la mayoría de estudios han estado orientados hacia los bosques húmedos tropicales. Los bosques secos se han reducido drásticamente y siguen muy amenazados, corriendo el riesgo de desaparecer en los próximos años. Por ello, es fundamental, generar investigación aplicada para la conservación inmediata de los ecosistemas secos tropicales. En Ecuador, la situación no es diferente y las zonas secas catalogadas como biodiversas están en constante amenaza. Los pocos estudios realizados en Ecuador sobre zonas secas, han permitido mejorar nuestro conocimiento referente a la diversidad y abundancia de las especies, relaciones planta-planta y síndromes de dispersión. No existen estudios sobre caracteres morfológicos en frutos y semillas de las especies leñosas de bosque seco. Sin embargo, nuestra comprensión de la dinámica y estructura de las comunidades ecológicas de zonas secas poco estudiadas, puede mejorar rápidamente mediante el estudio y enfoque de rasgos morfofisiológicos funcionales. El objetivo general del presente estudio fue aportar al conocimiento de la ecología y biología de semillas de zonas secas tropicales mediante el análisis y evaluación de rasgos morfofisiológicos de frutos y semillas de una comunidad de especies leñosas. El estudio se realizó en una zona de bosque y matorral seco, ubicados al sur occidente del Ecuador, a una altitud comprendida entre los 250 a 1 200 m s.n.m. caracterizada por una marcada estacionalidad ambiental, con lluvias desde diciembre a abril y una estación seca de mayo a noviembre. Precipitación media anual de 500 mm con una temperatura media anual de 20° a 26 °C. La zona de estudio forma parte de la región Tumbesina compartida entre el sur del Ecuador y el norte del Perú con gran diversidad de especies vegetales endémicas. Para el estudio se colectaron frutos con semillas maduras previamente a su dispersión de entre ocho y diez individuos de 80 especies entre árboles y arbustos más representativos de los bosques secos ecuatorianos. De los frutos colectados se utilizó una muestra al azar de 50 frutos y semillas por especie para los diferentes análisis. Se midió y evaluó 18 rasgos morfológicos y fisiológicos cuantitativos y cualitativos de frutos, semillas y de la especie. Se realizaron diferentes análisis de asociación y correlación entre los rasgos evaluados, con cinco variables ambientales registradas de las 109 parcelas establecidas en el área de estudio, además analizamos el tipo de dormición y comparamos la respuesta germinativa a la deshidratación relacionada con dos comunidades secas, matorral y bosque seco. Los resultados mostraron que las especies presentan gran heterogeneidad en rasgos continuos de las semillas. La variabilidad fue más evidente en rasgos como tamaño, volumen, masa y número de semillas por fruto. Sin embargo, una alta proporción de las especies tiende a producir una semilla por fruto. Además, la mayoría de las especies de bosque seco se caracterizan por no poseer algún tipo de apéndices o areola en sus semillas, forma ovalada y sin endospermo. La reserva nutritiva de las semillas se encuentra especialmente en los cotiledones de los embriones. Se encontraron seis tipos diferentes de embriones y la mayoría de las especies presentó embriones gruesos e invertidos. La dispersión de semillas está dominada por zoocoria en un 38 %, con relación a anemocoria (22 %) y autocoria (19 %). Sin embargo, encontramos que el 70 % de las especies posee frutos secos. Los análisis de dormición en las semillas de bosque seco, mostraron que el 60 % de las especies de bosque seco presentaron semillas con algún tipo de latencia, menor a la encontrada en especies de bosque deciduo tropical y sabanas, sin embargo, la dormición de las especies de bosque seco fue mayor al porcentaje de especies con dormición de bosque semiperenne y selva lluviosa tropical. La dormición física constituyó el 35 % de las especies de bosque seco, seguido del 12 % con dormición fisiológica, mientras que solamente una especie tuvo dormición morfológica. Encontramos que la dormición de las semillas de las especies en estudio se relaciona significativamente con el tipo y función del embrión y con el endospermo. Existieron relaciones significativas entre los rasgos morfológicos de los frutos, semillas, embriones y atributos de los individuos de 46 especies, aunque en algunos casos con coeficientes de correlación bajos. Hubo pocas relaciones entre los rasgo morfológicos de las semillas con las variables ambientales registradas. Solamente el tipo de testa y la presencia de apéndices en las semillas mostraron relación con el pH y la temperatura media del suelo. No obstante usando el modelo fouth corner-RLQ, no se encontraron asociaciones claras ni significativas entre rasgos morfológicos de semillas y frutos con variables ambientales. Al medir el efecto de la deshidratación en las semillas de los dos hábitats secos tropicales: bosque y matorral seco, los resultados determinaron que tanto las semillas de las especies leñosas de ambientes más áridos (matorral seco) están en gran medida pre-adaptadas a la desecación que las especies de ambientes menos áridos (bosque seco). Los tratamientos de deshidratación ejercieron un efecto negativo en los porcentajes de germinación en todas las especies, excepto para C. platanifolia. Los resultados más sorprendentes se registraron para Senna alata que mostró germinación extremadamente baja o incluso sin germinación a contenidos de humedad de la semillas de 0,10 g H2O g de peso seco. Las curvas de germinación difirieron significativamente entre los tratamientos de deshidratación en cada especie. Aportar al conocimiento la fisiología de la deshidratación y los límites de tolerancia de las semillas de bosque y matorral seco ayudará a entender mejor el papel de este rasgo en la ecología de las semillas y dinámica de las comunidades áridas tropicales. El estudio demostró, que la adaptación ecológica de las semillas de las especies leñosas de bosque seco a factores ambientales extremos, puede verse reflejada en una red de interacciones y correlaciones complejas entre los propios rasgos morfológicos y fisiológicos continuos y cuantitativos, sobre todo en rasgos internos de las semillas, quienes ejercerían una mayor influencia en toda la red de interacciones. Si bien, los rasgos de las semillas no mostraron fuertes relaciones con las variables ambientales, posiblemente las asociaciones presentes entre rasgos morfológicos pudiesen predecir en cambio interacciones entre especies y comportamientos y procesos relacionados con la tolerancia a la deshidratación y dormición de las semillas. ABSTRACT South America is one of the areas with the largest number of tropical dry forest in the world. However, these forests have been poorly understood and most studies have been directed to tropical rainforests. Dry forests have been drastically reduced and are very threatened, risking desaparecerer in the next years. It is therefore essential, generate applied research for conservation of tropical dry ecosystems. In Ecuador the situation is no different and dry areas classified as biodiverse are under constant threat. The few studies made in Ecuador on drylands have improved our knowledge concerning the diversity and abundance of species, plant-plant relationships and dispersion syndromes. Morphological studies on fruits and seeds of woody dry forest species do not exist. However, our understanding of the dynamics and structure of ecological communities dryland little studied, may improve quickly through the study and functional approach morphophysiological traits. The overall objective of this study was to contribute to the knowledge of the ecology and biology of tropical dry seeds through analysis and evaluation of morphophysiological traits of fruits and seeds of a community of woody species. The study was conducted in an area of dry scrub forest, located at the southwest of Ecuador, at an altitude between 250 to 1200 m asl. Environmental characterized by a marked seasonality, with rainfall from December to April and a dry season from May to November. Annual rainfall of 500 mm with an average annual temperature of 20° to 26 °C. The study area is part of the shared Tumbesina region between southern Ecuador and northern Peru with a great diversity of endemic plant species. For the study, we collected fruit and seed madure of eight and ten individuos of 80 species of trees and shrub most representated of the Ecuador dry forest. We selected a sample of 50 fruits and seeds for different analysis. We measure and evaluate 18 morphological and physiological traits of fruits, seeds and species. We perform analysis and correlation between traits associated with five environmental variables taken from the 109 plots established in the study area also analyze and compare the germination response to dehydration related to two dry communities, scrub and dry forest. The results showed that the species have great heterogeneity in continuous seed traits. Variability was more evident in features such as size, volume, mass, and number of seeds per fruit. However, a high proportion of species tends to produce a seed per fruit. In addition, most of the species of dry forest is characterized by not having some sort of ppendices or areola in its seeds, oval form and without endosperm. The nutrient reserves of seeds are especially in the cotyledons of the embryos. Six different embryos were found and most of the species presented thick and inverted embryos. Seed dispersal zoochory is dominated by 38 %, relative to anemochory (22 %) and autochory (19 %). However, we found that 70 % of the species has dried fruits. The analysis of dormancy from tropical dry forest, showed that 60 % of species showed seed dormancy, down from species found in tropical deciduous forest and savanna, however dormancy dry forest species was higher than the percentage of forest species dormancy semi-evergreen and tropical rain forest. Physical dormancy corresponds to 35 % of species, followed by 12 % with physiological dormancy, while only one species had morphological dormancy. We found that dormancy of the seeds was significantly related to the type and function of the embryo and the endospemo. There were significant relationships between morphological traits of fruits, seeds, embryos and attributes of individuals of 46 species, although in some cases with low correlation coefficients. There was little relationship between the morphologic traits of the seeds with the registered environmental variables. Only the type of tesla and the presence of appendages on the seeds showed relation to pH and the mean soil temperature. However, using the fourth corner-RLQ model, neither clear nor significant between morphological traits of seeds and fruits associations with environmental variables were found. The effect of dehydration on seeds of two tropical dry forest habitats was evident in dry scrub. The results determined that both the seeds of woody species forest and dry scrub are pre-adapted to drier conditions. Dehydration treatments exerted a negative effect on germination percentage in all species, except for C. platanifolia. However, all species germinated in treatments of extreme dryness, but in low percentages. The most striking results were recorded for Senna alata showed no germination when its moisture content was 0.10 g H2O g dry weight. Germination curves differ significantly between the treatments of dehydration in each species. Contribute to the knowledge of physiology and dehydration tolerance limits seeds dry scrub forest and help you better understand the role of this trait in seed ecology and dynamics of tropical arid communities. The study showed that the ecological adaptation of seeds of woody species of dry forest to extreme environmental factors may be reflected in a complex web of interactions and correlations between morphological and physiological traits continuous and quantitative themselves, especially in internal seed traits, who exerted a major influence on the entire network of interactions. While the seed traits showed strong relationships with environmental variables possibly present associations between morphological traits could predict interactions between species and change behaviors related to desiccation tolerance and seed dormancy processes.

Different response to Plasmodium falciparum malaria in West African sympatric ethnic groups

The comparison of malaria indicators among populations that have different genetic backgrounds and are uniformly exposed to the same parasite strains is one approach to the study of human heterogeneities in the response to the infection. We report the results of comparative surveys on three sympatric West African ethnic groups, Fulani, Mossi, and Rimaibé, living in the same conditions of hyperendemic transmission in a Sudan savanna area northeast of Ouagadougou, Burkina Faso. The Mossi and Rimaibé are Sudanese negroid populations with a long tradition of sedentary farming, while the Fulani are nomadic pastoralists, partly settled and characterized by non-negroid features of possible caucasoid origin. Parasitological, clinical, and immunological investigations showed consistent interethnic differences in Plasmodium falciparum infection rates, malaria morbidity, and prevalence and levels of antibodies to various P. falciparum antigens. The data point to a remarkably similar response to malaria in the Mossi and Rimaibé, while the Fulani are clearly less parasitized, less affected by the disease, and more responsive to all antigens tested. No difference in the use of malaria protective measures was demonstrated that could account for these findings, and sociocultural or environmental factors do not seem to be involved. Known genetic factors of resistance to malaria did not show higher frequencies in the Fulani. The differences in the immune response were not explained by the entomological observations, which indicated substantially uniform exposure to infective bites. The available data support the existence of unknown genetic factors, possibly related to humoral immune responses, determining interethnic differences in the susceptibility to malaria.

Fatores abióticos condicionantes da distribuição de espécies arbóreas em quatro formações florestais do Estado de São Paulo

No estudo das comunidades florestais, estabelecer a importância relativa dos fatores que definem a composição e a distribuição das espécies é um desafio. Em termos de gradientes ambientais o estudo das respostas das espécies arbóreas são essenciais para a compreensão dos processos ecológicos e decisões de conservação. Neste sentido, para contribuir com a elucidação dos processos ecológicos nas principais formações florestais do Estado de São Paulo (Floresta Ombrófila Densa de Terras Baixas, Floresta Ombrófila Densa Submontana, Floresta Estacional Semidecidual e Savana Florestada) este trabalho objetivou responder as seguintes questões: (I) a composição florística e a abundância das espécies arbóreas, em cada unidade fitogeográfica, variam conforme o gradiente edáfico e topográfico?; (II) características do solo e topografia podem influenciar na previsibilidade de ocorrência de espécies arbóreas de ampla distribuição em diferentes tipos vegetacionais? (III) existe relação entre o padrão de distribuição espacial de espécies arbóreas e os parâmetros do solo e topografia? O trabalho foi realizado em parcelas alocadas em unidades de conservação (UC) que apresentaram trechos representativos, em termos de conservação e tamanho, das quatro principais formações florestais presentes no Estado de São Paulo. Em cada UC foram contabilizados os indivíduos arbóreos (CAP ≥ 15 cm), topografia, dados de textura e atributos químicos dos solos em uma parcela de 10,24 ha, subdividida em 256 subparcelas. Análises de correspodência canônica foram aplicadas para estabelecer a correspondência entre a abundância das espécies e o gradiente ambiental (solo e topografia). O método TWINSPAN modificado foi aplicado ao diagrama de ordenação da CCA para avaliar a influência das variáveis ambientais (solo e topografia) na composição de espécies. Árvores de regressão \"ampliadas\" (BRT) foram ajustadas para a predição da ocorrência das espécies segundo as variáveis de solo e topografia. O índice de Getis-Ord (G) foi utilizado para determinar a autocorrelação espacial das variáveis ambientais utilizadas nos modelos de predição da ocorrência das espécies. Nas unidades fitogeográficas analisadas, a correspondência entre o gradiente ambiental (solo e topografia) e a abundância das espécies foi significativa, especialmente na Savana Florestada onde observou-se a maior relação. O solo e a topografia também se relacionaram com a semelhança na composição florística das subparcelas, com exceção da Floresta Estacional Semicidual (EEC). As principais variáveis de solo e topografia relacionadas a flora em cada UC foram: (1) Na Floresta Ombrófila Densa de Terras Baixas (PEIC) - teor de alumínio na camada profunda (Al (80-100 cm)) que pode refletir os teor de Al na superfície, acidez do solo (pH(H2O) (5-25 cm)) e altitude, que delimitou as áreas alagadas; (2) Na Floresta Ombrófila Densa Submontana (PECB) - altitude, fator que, devido ao relevo acidentado, influencia a temperatura e incidência de sol no sub-bosque; (3) Na Savana Florestada (EEA) - fertilidade, tolerância ao alumínio e acidez do solo. Nos modelos de predição BRT, as variáveis químicas dos solos foram mais importantes do que a textura, devido à pequena variação deste atributo no solo nas áreas amostradas. Dentre as variáveis químicas dos solos, a capacidade de troca catiônica foi utilizada para prever a ocorrência das espécies nas quatro formações florestais, sendo particularmente importante na camada mais profunda do solo da Floresta Ombrófila Densa de Terras Baixas (PEIC). Quanto à topografia, a altitude foi inserida na maioria dos modelos e apresentou diferentes influências sobre as áreas de estudo. De modo geral, para presença das espécies de ampla distribuição observou-se uma mesma tendência quando à associação com os atributos dos solos, porém com amplitudes dos descritores edáficos que variaram de acordo com a área de estudo. A ocorrência de Guapira opposita e Syagrus romanzoffiana, cujo padrão variou conforme a escala, foi explicada por variáveis com padrões espaciais agregados que somaram entre 30% e 50% de importância relativa no modelo BRT. A presença de A. anthelmia, cujo padrão também apresentou certo nível de agregação, foi associada apenas a uma variável com padrão agregado, a altitude (21%), que pode ter exercido grande influência na distribuição da espécie ao delimitar áreas alagadas. T. guianensis se associou a variáveis ambientais preditoras com padrão espacial agregado que somaram cerca de 70% de importância relativa, o que deve ter sido suficiente para estabelecer o padrão agregado em todas as escalas. No entanto, a influência dos fatores ambientais no padrão de distribuição da espécie não depende apenas do ótimo ambiental da espécie, mas um resultado da interação espécie-ambiente. Concluiu-se que: (I) características edáficas e topográficas explicaram uma pequena parcela da composição florística, em cada unidade fitogeográfica, embora a ocorrência de algumas espécies tenha se associado ao gradiente edáfico e topográfico; (II) a partir de características dos solos e da topografia foi possível prever a presença de espécies arbóreas, que apresentaram particularidades em relação a sua associação com o solo de cada fitofisionomia; (III) a partir de associações descritivas o solo e a topografia influenciam o padrão de distribuição espacial das espécies, na proporção em que contribuem para a presença das mesmas.

The late Pliocene pollen record of ODP Site 108-658 off Cape Blanc, Atlantic Ocean

A 200 m long marine pollen record from ODP Site 658 (21°N, 19°W) reveals cyclic fluctuations in vegetation and continental climate in northwestern Africa from 3.7 to 1.7 Ma. These cycles parallel oxygen isotope stages. Prior to 3.5 Ma, the distribution of tropical forests and mangrove swamps reached Cape Blanc, 5°N of the present distribution. Between 3.5 and 2.6 Ma, forests occurred at this latitude during irregular intervals and nearly disappeared afterwards. Likewise, a Saharan paleoriver flowed continuously until isotope Stage 134 (3.35 Ma). When river discharge ceased, wind transport of pollen grains prevailed over fluvial transport. Pollen indicators of trade winds gradually increased between 3.3 and 2.5 Ma. A strong aridification of the climate of northwestern Africa occurred during isotope Stage 130 (3.26 Ma). Afterwards, humid conditions reestablised followed by another aridification around 2.7 Ma. Repetitive latitudinal shifts of vegetation zones ranging from wooded savanna to desert flora dominated for the first time between between 2.6 and 2.4 Ma as a response to the glacial stages 104, 100 and 98. Although climatic conditions, recorded in the Pliocene, were not as dry as those of the middle and Late Pleistocene, latitudinal vegetation shifts near the end of the Pliocene resembled those of the interglacial-glacial cycles of the Brunhes chron.

Changes in the vegetation and trade winds in equatorial Northwest Africa 140,000-70,000 yr B.P., as deduced from two marine pollen records

The deep-sea cores M 16415-2 and M 16416-2 at about 9°N off Sierra Leone were analysed palynologically for the time interval 140,000-70,000 yr B.P. Results were presented in absolute (pollen concentration and pollen influx) and relative diagrams (pollen percentage). In a previous study it was evidenced that in northwest Africa pollen is mainly transported to the Atlantic by wind, so that the efficiency of aeolian pollen transport (pollen flux) could be used to evaluate changes in the intensity of the northeast trade winds. The glacial episodes (represented by the oxygen isotope stages 6 and 4) are characterized by strong northeast trade winds, whereas the last interglacial (stage 5) is characterized by weak trade winds. The pollen influx diagram shows that the intensity of the trade winds increased slightly during the relatively cool intervals of stage 5 (viz. 5.4 and 5.2). Tropical forest had maximally expanded around 124,000 yr B.P. (stage 5.5), around 98,000 yr B.P. (transition of stage 5.3 to 5.2), and around 70,000 yr B.P. (first part of stage 4): an increasing delay of the response of tropical forest to global intervals with maximum temperature is apparent during the last interglacial. As tropical forests need continuous humidity, the record of tropical forest monitors changes in climatic humidity south of the Sahara. During the last interglacial, the southern boundary of the Sahara shifted only little: expansions and contractions of the tropical forest area are correlated with contra-oscillations of the grass-dominated savanna zone. Great latitudinal shifts of the desert savanna boundary, on the contrary, occurred during the penultimate glacial interglacial transition (around 128,000 yr B.P.) to the north, and during the last interglacial-glacial transition (around 65,000 yr B.P.) to the south.

Global 2009 1km MODIS (MOD35/MOD09) cloud frequency and MOD35 processing path

Identifying cloud interference in satellite-derived data is a critical step toward developing useful remotely sensed products. Most MODIS land products use a combination of the MODIS (MOD35) cloud mask and the 'internal' cloud mask of the surface reflectance product (MOD09) to mask clouds, but there has been little discussion of how these masks differ globally. We calculated global mean cloud frequency for both products, for 2009, and found that inflated proportions of observations were flagged as cloudy in the Collection 5 MOD35 product. These erroneously categorized areas were spatially and environmentally non-random and usually occurred over high-albedo land-cover types (such as grassland and savanna) in several regions around the world. Additionally, we found that spatial variability in the processing path applied in the Collection 5 MOD35 algorithm affects the likelihood of a cloudy observation by up to 20% in some areas. These factors result in abrupt transitions in recorded cloud frequency across landcover and processing-path boundaries impeding their use for fine-scale spatially contiguous modeling applications. We show that together, these artifacts have resulted in significantly decreased and spatially biased data availability for Collection 5 MOD35-derived composite MODIS land products such as land surface temperature (MOD11) and net primary productivity (MOD17). Finally, we compare our results to mean cloud frequency in the new Collection 6 MOD35 product, and find that landcover artifacts have been reduced but not eliminated. Collection 6 thus increases data availability for some regions and land cover types in MOD35-derived products but practitioners need to consider how the remaining artifacts might affect their analysis.

Table 1: Main characteristics of the BIOTA observatories included in this study & Table 3: Occurrence of termite taxa at the observatories

Termites are the most important soil ecosystem engineers of semi-arid and arid habitats. They enhance decomposition processes as well as the subsequent mineralisation of nutrients by bacteria and fungi. Through their construction of galleries, nests and mounds, they promote soil turnover and influence the distribution of nutrients and also alter texture and hydrological properties of soils, thereby affecting the heterogeneity of their ecosystem. The main aim of the present thesis was to define the impact of termites on ecosys-tem functioning in a semi-arid ecosystem. In a baseline study, I assessed the diversity of termite taxa in relation to the amount of precipitation, the vegetation patterns and the land use systems at several sites in Namibia. Subsequently, I focussed on a species that is highly abundant in many African savannas, the fungus growing and mound building species Macro-termes michaelseni (Sjöstedt, 1914). I asked how this species influences the spatial hetero-geneity of soil and vegetation patterns. From repeated samplings at 13 sites in Namibia, I obtained 17 termite taxa of 15 genera. While the type of land use seems to have a minor effect on the termite fauna, the mean annual precipitation explained 96% and the Simpson index of vascular plant diversity 81% of the variation in taxa diversity. The number of termite taxa increased with both of these explanation variables. In contrast to former studies on Macrotermes mounds in several regions of Africa that I reviewed, soil analyses from M. michaelseni mounds in the central Namibian savanna revealed that they contain much higher nitrogen contents when compared to their parent material. Further analyses revealed that nitrate forms a major component of the nitrogen content in termite mounds. As nitrate solves easily in water, evaporation processes are most probably responsible for the transport of solved nitrates to the mound surface and their accumulation there. The analysed mounds in central Namibia contained higher sand propor-tions compared to the mounds of the former studies. Through the higher percentage of coarse and middle sized pores, water moves more easily in sandy soils compared to more clayey soils. In consequence, evaporation-driven nitrate accumulation can occur in the studied mounds at high rates. ff...

(Fig. 9) Pollen analysis of sediment core GIK16017-2

The improved understanding of the pollen signal in the marine sediments offshore of northwest Africa is applied to deep-sea core M 16017-2 at 21°N. Downcore fluctuations in the percentage, concentration and influx diagrams record latitudinal shifts of the main northwest African vegetation zones and characteristics of the trade winds and the African Easterly Jet. Time control is provided by 14C ages and 180 records. During the period 19,000-14,000 yr B.P. a compressed savanna belt extended between about 12 ° and 14-15°N. The Sahara had maximally expanded northward and southward under hyperarid climatic conditions. The belt with trade winds and dominant African Easterly Jet transport had not shifted latitudinally. The trade winds were strong as compared to the modern situation but around 13,000 yr B.P. the trade winds weakened. After 14,000 yr B.P. the climate became less arid south of the Sahara and a first spike of fluvial runoff is registered around 13,000 yr B.P. Fluvial runoff increased strongly around 11,000 yr B.P. and maximum runoff is recorded from about 9000-7800 yr B.P. Around 12,500 yr B.P. the savanna belt started to shift northward and became richer in woody species: it shifted about 6° of latitude, reached its northernmost position during the period of 9200-7800 yr B.P. and extended between about 16° and 24°N at that time. Tropical forest had reached its maximum expansion and the Guinea zone reached as far north as about 15°N, reflecting very humid climatic conditions south of the Sahara. North of the Sahara the climate also became more humid and Mediterranean vegetation developed rapidly. The Sahara had maximally contracted and the trade winds were weak and comparable with the present day intensity. After about 7800 yr B.P. the southern fringe of the Sahara and accordingly the savanna belt, shifted rapidly southward again.

Das illustrirte Mississippithal : dargestellt in 80 nach der Natur aufgenommenen Ansichten vom Wasserfalle zu St. Anthony an bis zum Golf von Mexico (eine Entfernung von ungefähr 2300 englischen Meilen) /

Cf. Sabin 40807.

Hazard classification of United States military explosives and hazardous munitions.

Cover title.

Adaptations of strangler figs to life in the rainforest canopy

Figs are rainforest keystone species. Non-strangler figs establish on the forest floor; strangler figs establish epiphytically, followed by a dramatic transition from epiphyte to free-standing tree that kills its hosts. Free-standing figs display vigorous growth and resource demand suggesting that epiphytic strangler figs require special adaptations to deal with resource limitations imposed by the epiphytic environment. We studied epiphytic and free-standing strangler figs, and non-strangler figs in tropical rainforest and in cultivation, as well as strangler figs in controlled conditions. We investigated whether the transition from epiphyte to free-standing tree is characterised by morphological and physiological plasticity. Epiphyte substrate had higher levels of plant-available ammonium and phosphate, and similar levels of nitrate compared with rainforest soil, suggesting that N and P are initially not limiting resources. A relationship was found between taxonomic groups and plant N physiology; strangler figs, all members of subgenus Urostigma, had mostly low foliar nitrate assimilation rates whereas non-strangler figs, in subgenera Pharmacocycea, Sycidium, Sycomorus or Synoecia, had moderate to high rates. Nitrate is an energetically expensive N source, and low nitrate use may be an adaptation of strangler figs for conserving energy during epiphytic growth. Interestingly, significant amounts of nitrate were stored in fleshy taproot tubers of epiphytic stranglers. Supporting the concept of plasticity, leaves of epiphytic Ficus benjamina L. had lower N and C content per unit leaf area, lower stomatal density and 80% greater specific leaf area than leaves of conspecific free-standing trees. Similarly, glasshouse-grown stranglers strongly increased biomass allocation to roots under water limitation. Epiphytic and free-standing F. benjamina had similar average foliar delta C-13, but epiphytes had more extreme values; this indicates that both groups of plants use the C-3 pathway of CO2 fixation but that water availability is highly variable for epiphytes. We hypothesise that epiphytic figs use fleshy stem tubers to avoid water stress, and that nitrate acts as an osmotic compound in tubers. We conclude that strangler figs are a unique experimental system for studying the transition from rainforest epiphyte to tree, and the genetic and environmental triggers involved.