Global beta-diversity of angiosperm trees is shaped by Quaternary climate change.

As Earth's climate has varied strongly through geological time, studying the impacts of past climate change on biodiversity helps to understand the risks from future climate change. However, it remains unclear how paleoclimate shapes spatial variation in biodiversity. Here, we assessed the influence of Quaternary climate change on spatial dissimilarity in taxonomic, phylogenetic, and functional composition among neighboring 200-kilometer cells (beta-diversity) for angiosperm trees worldwide. We found that larger glacial-interglacial temperature change was strongly associated with lower spatial turnover (species replacements) and higher nestedness (richness changes) components of beta-diversity across all three biodiversity facets. Moreover, phylogenetic and functional turnover was lower and nestedness higher than random expectations based on taxonomic beta-diversity in regions that experienced large temperature change, reflecting phylogenetically and functionally selective processes in species replacement, extinction, and colonization during glacial-interglacial oscillations. Our results suggest that future human-driven climate change could cause local homogenization and reduction in taxonomic, phylogenetic, and functional diversity of angiosperm trees worldwide.

A potential consequence for urban birds' fitness: Exposed anthropogenic nest materials reduce nest survival in the clay-colored thrush.

Urban bird dwellers survive and reproduce in highly urbanized ecosystems. Some individuals adjust to these novel conditions by changing natural nesting materials for artificial materials, thus making nests more conspicuous in the environment. The consequences of using artificial materials for nesting remain poorly understood, especially from a nest-predator perspective. We studied if exposed artificial materials on bird nests affect the daily survival rate in a common dweller species, the clay-colored thrush (Turdus grayi). We used previously collected nests with a different area of exposed artificial materials and placed them with clay eggs on the main campus of the Universidad de Costa Rica. During 12 days of the reproductive season, we monitored the nests using trap cameras located in front of each nest. We found that nest survival decreased as the proportion of exposed artificial materials increased in the nest and, unexpectedly, that the principal predators were conspecifics. Thus, artificial materials used in the outer layer of nests make them more susceptible to predation. The use of artificial materials likely reduce reproductive success and population size in urban clay-colored thrush, but further field experiments measuring the effect of waste in birds' nests on the reproductive success of urban birds are needed.

High exposure of global tree diversity to human pressure.

Safeguarding Earth's tree diversity is a conservation priority due to the importance of trees for biodiversity and ecosystem functions and services such as carbon sequestration. Here, we improve the foundation for effective conservation of global tree diversity by analyzing a recently developed database of tree species covering 46,752 species. We quantify range protection and anthropogenic pressures for each species and develop conservation priorities across taxonomic, phylogenetic, and functional diversity dimensions. We also assess the effectiveness of several influential proposed conservation prioritization frameworks to protect the top 17% and top 50% of tree priority areas. We find that an average of 50.2% of a tree species' range occurs in 110-km grid cells without any protected areas (PAs), with 6,377 small-range tree species fully unprotected, and that 83% of tree species experience nonnegligible human pressure across their range on average. Protecting high-priority areas for the top 17% and 50% priority thresholds would increase the average protected proportion of each tree species' range to 65.5% and 82.6%, respectively, leaving many fewer species (2,151 and 2,010) completely unprotected. The priority areas identified for trees match well to the Global 200 Ecoregions framework, revealing that priority areas for trees would in large part also optimize protection for terrestrial biodiversity overall. Based on range estimates for >46,000 tree species, our findings show that a large proportion of tree species receive limited protection by current PAs and are under substantial human pressure. Improved protection of biodiversity overall would also strongly benefit global tree diversity.

The predictive power of pollination syndromes: Passerine pollination in heterantherous Meriania macrophylla (Benth.) Triana (Melastomataceae).

The cloud forest species Meriania macrophylla (Benth.) Triana has pseudocampanulate flowers with bulbous stamen appendages, typical for the passerine pollination syndrome found in the Melastomataceae tribe Merianieae. The species is further characterized by strong stamen dimorphism (heteranthery), a condition otherwise associated with pollen-rewarding bee-pollinated species (both in Melastomataceae and beyond). In passerine-pollinated Merianieae, however, flowers usually only show weak stamen dimorphism. Here, we conducted field and laboratory investigations to determine the pollinators of M. macrophylla and assess the potential role of strong heteranthery in this species. Our field observations in Costa Rica confirmed syndrome predictions and indeed proved pollination by passerine birds in M. macrophylla. The large bulbous set of stamens functions as a food-body reward to the pollinating birds, and as trigger for pollen release (bellows mechanism) as typical for the passerine syndrome in Merianieae. In contrast to other passerine-pollinated Merianieae, the second set of stamens has seemingly lost its rewarding and pollination function, however. Our results demonstrate the utility of the pollination syndrome concept even in light of potentially misleading traits such as strong heteranthery.

La especie de bosque nuboso Meriania macrophylla (Benth.) Triana presenta flores pseudocampanuladas con estambres con un apéndice bulboso, una morfología típica de las flores polinizadas por aves paserinas en Merianieae. Además, la especie se caracteriza por tener dos verticilos diferentes de estambres (heteroanteria), una condición que generalmente se asocia a especies polinizadas por abejas con polen como recompensa. En las especies de Merianieae polinizadas por aves la heteroantería es leve, si presente. Para este estudio determinamos los polinizadores de M. macrophylla y evaluamos posibles explicaciones de la presencia de estambres heteroantéricos Nuestras observaciones de campo en Costa Rica confimaron nuestras predicciones de que la polinización de esta especie se da por aves. Los conectivos engrosados funcionan como recompensa para las aves, que al removerlos de la flor activan el mecanismo de expulsión de polen. En contraste, el segundo verticilo de estambres aparentemente perdió tanto la función de recompensa como su función reproductiva. Nuestros resultados demuestran la utilidad de los síndromes de polinización incluso en presencia de caracteres confusos como la presencia de heteroanteria muy marcada.

Human waste used as nesting material affects nest cooling in the clay-colored thrush.

The internal temperature of nests largely depends on the materials used in their construction because the characteristics of each material affect the isolation of nest walls. In urban environments, the availability of natural materials for nest building decreases, while the availability of artificial materials increases. Therefore, many urban bird species use more artificial materials for nest building inside cities, which may affect the thermal properties of the nest. We conducted an experiment to measure the effect of artificial materials included as part of the nest structure, on nest thermoregulation. We used as a model, nests of the clay-colored thrush (Turdus grayi), an urban bird species that have been reported using artificial nest materials. In our experiment, we measured how variation in artificial materials mass affects the nest cooling rate in a climate-controlled room. We found that artificial materials increased the cooling rate of clay-colored thrush nests, compared with nests with only natural materials. This result is especially relevant because showed a negative direct effect of the use of artificial material for nest building in birds. Considering that the availability of artificial material is increasing in urban areas, while natural material is decreasing, it is expected that the negative effect of using artificital material for nest construction would increase in the clay-colored thrush and other city bird species.

Dimensions of invasiveness: Links between local abundance, geographic range size, and habitat breadth in Europe's alien and native floras.

Understanding drivers of success for alien species can inform on potential future invasions. Recent conceptual advances highlight that species may achieve invasiveness via performance along at least three distinct dimensions: 1) local abundance, 2) geographic range size, and 3) habitat breadth in naturalized distributions. Associations among these dimensions and the factors that determine success in each have yet to be assessed at large geographic scales. Here, we combine data from over one million vegetation plots covering the extent of Europe and its habitat diversity with databases on species' distributions, traits, and historical origins to provide a comprehensive assessment of invasiveness dimensions for the European alien seed plant flora. Invasiveness dimensions are linked in alien distributions, leading to a continuum from overall poor invaders to super invaders-abundant, widespread aliens that invade diverse habitats. This pattern echoes relationships among analogous dimensions measured for native European species. Success along invasiveness dimensions was associated with details of alien species' introduction histories: earlier introduction dates were positively associated with all three dimensions, and consistent with theory-based expectations, species originating from other continents, particularly acquisitive growth strategists, were among the most successful invaders in Europe. Despite general correlations among invasiveness dimensions, we identified habitats and traits associated with atypical patterns of success in only one or two dimensions-for example, the role of disturbed habitats in facilitating widespread specialists. We conclude that considering invasiveness within a multidimensional framework can provide insights into invasion processes while also informing general understanding of the dynamics of species distributions.

Mechanisms driving plant functional trait variation in a tropical forest.

Plant functional trait variation in tropical forests results from taxonomic differences in phylogeny and associated genetic differences, as well as, phenotypic plastic responses to the environment. Accounting for the underlying mechanisms driving plant functional trait variation is important for understanding the potential rate of change of ecosystems since trait acclimation via phenotypic plasticity is very fast compared to shifts in community composition and genetic adaptation. We here applied a statistical technique to decompose the relative roles of phenotypic plasticity, genetic adaptation, and phylogenetic constraints. We examined typically obtained plant functional traits, such as wood density, plant height, specific leaf area, leaf area, leaf thickness, leaf dry mass content, leaf nitrogen content, and leaf phosphorus content. We assumed that genetic differences in plant functional traits between species and genotypes increase with environmental heterogeneity and geographic distance, whereas trait variation due to plastic acclimation to the local environment is independent of spatial distance between sampling sites. Results suggest that most of the observed trait variation could not be explained by the measured environmental variables, thus indicating a limited potential to predict individual plant traits from commonly assessed parameters. However, we found a difference in the response of plant functional traits, such that leaf traits varied in response to canopy-light regime and nutrient availability, whereas wood traits were related to topoedaphic factors and water availability. Our analysis furthermore revealed differences in the functional response of coexisting neotropical tree species, which suggests that endemic species with conservative ecological strategies might be especially prone to competitive exclusion under projected climate change.

Climatic and edaphic controls over tropical forest diversity and vegetation carbon storage.

Tropical rainforests harbor exceptionally high biodiversity and store large amounts of carbon in vegetation biomass. However, regional variation in plant species richness and vegetation carbon stock can be substantial, and may be related to the heterogeneity of topoedaphic properties. Therefore, aboveground vegetation carbon storage typically differs between geographic forest regions in association with the locally dominant plant functional group. A better understanding of the underlying factors controlling tropical forest diversity and vegetation carbon storage could be critical for predicting tropical carbon sink strength in response to projected climate change. Based on regionally replicated 1-ha forest inventory plots established in a region of high geomorphological heterogeneity we investigated how climatic and edaphic factors affect tropical forest diversity and vegetation carbon storage. Plant species richness (of all living stems >10 cm in diameter) ranged from 69 to 127 ha-1 and vegetation carbon storage ranged from 114 to 200 t ha-1. While plant species richness was controlled by climate and soil water availability, vegetation carbon storage was strongly related to wood density and soil phosphorus availability. Results suggest that local heterogeneity in resource availability and plant functional composition should be considered to improve projections of tropical forest ecosystem functioning under future scenarios.

Concentración foliar de macro- y micro-nutrimentos en cuatro leguminosas maderables del trópico estacionalmente seco de Costa Rica / Foliar concentration of macro- and micronutrients in four woody legumes from a tropical dry-seasonal forest in Costa Rica

Resumen La productividad de las plantaciones forestales con especies nativas se puede incrementar si se conocen bien los requerimientos nutricionales de cada una de las especies. Estos requerimientos pueden variar de acuerdo a las estrategias ecológicas de las especies y a las adaptaciones que tienen, como por ejemplo la capacidad de fijar nitrógeno. En el presente trabajo se presentan los "rangos satisfactorios" de concentración foliar de macronutrimentos y micronutrimentos para cuatro leguminosas del trópico estacionalmente seco de Guanacaste, Costa Rica. Se analizó la concentración foliar de N, P, Ca, Mg, K, S, Fe, Mn, Cu, Zn y B, Na en las especies Dalbergia retusa (cocobolo), Enterolobium cyclocarpum (guanacaste), Hymenaea courbaril (guapinol) y Schizolobium parahyba (gallinazo). La concentración de nutrimentos varió con las especies y según la capacidad de fijar nitrógeno o no y dentro de cada una de esas categorías de acuerdo a la tasa de crecimiento. La concentración total de nutrimentos fue mayor para las especies fijadoras de N que para las no fijadoras y dentro de cada uno de esos grupos la concentración total de nutrimentos fue mayor para las especies de crecimiento rápido comparado con aquellas de crecimiento más lento. De acuerdo a la cantidad de nutrimentos observados en las hojas, se recomienda que H. courbaril y D. retusa deberían ser sembradas en suelos relativamente pobres de nutrimentos, mientras que E. cyclocarpum y S. parahyba en suelos más fértiles. Para D. retusa e H. courbaril se presenta información nueva sobre concentraciones y se mejora la misma para algunos elementos no mencionados en la literatura para E. cyclocarpum y S. parahyba.

Abstract The information about nutritional aspects of trees in the tropical dry forest is scarce. This investigation aims to establish a normal range of foliar nutrient concentration to four forestry species as related to their growth rates and biological nitrogen fixation capacity in the seasonally dry ecosystem at Guanacaste, Costa Rica. Foliar samples for four species: Dalbergia retusa (cocobolo), Enterolobium cyclocarpum (guanacaste), Hymenaea courbaril (guapinol or jatobá) and Schizolobium parahyba (gallinazo or pachaco), all belonging to the Fabaceae family, were analyzed for N, P, Ca, Mg, K, S, Fe, Mn, Cu, Zn, B and Na. Nutrient concentration varied with species according to their ability to fix nitrogen and their rate of growth. The overall concentration of macronutrients was higher in nitrogen-fixing species compared with non-fixing species, and within each one of those groups, the species with higher growth rate presented more macronutrients than species with low growth rate. According to the concentration of foliar nutrients accumulated in the foliage, we recommend that H. courbaril and D. retusacould be grown in the region soils with medium fertility and E. cyclocarpum and S. parahyba in the more fertile soils. Values ranges of foliar concentrations considered as satisfactory presented in this work are new for D. retusa and H. courbaril and improve the scarce information available for E. cyclocarpum and S. parahyba. Rev. Biol. Trop. 66(3): 969-983. Epub 2018 September 01.

Is local trait variation related to total range size of tropical trees?

The reasons why the range size of closely related species often varies significantly have intrigued scientists for many years. Among other hypotheses, species with high trait variation were suggested to occupy more diverse environments, have more continuity in their distributions, and consequently have larger range sizes. Here, using 34 tree species of lowlands tropical rainforest in southern Costa Rica, we explored whether inherent trait variability expressed at the local scale in functional traits is related to the species' total geographical range size. We formed 17 congeneric pairs of one narrow endemic and one widespread species, sampled 335 individuals and measured eight functional traits: leaf area, leaf thickness, leaf dry matter content, specific leaf area, leaf nitrogen content, leaf phosphorus content, leaf nitrogen to phosphorus ratio, and wood specific gravity. We tested whether there are significant differences in the locally expressed variation of individual traits or in multidimensional trait variance between the species in congeneric pairs and whether species' range size could hence be predicted from local trait variability. However, we could not find such differences between widely distributed and narrow range species. We discuss the possible reasons for these findings including the fact that higher trait variability of widespread species may result from successive local adaptations during range expansion and may hence often be an effect rather than the cause of larger ranges.

Seedling growth and survival of five tree species in secondary forests and adjacent pastures in the montane rain forests of Southern Costa Rica / Crecimiento y supervivencia de plántulas de cinco especies de árboles en bosques secundarios y pastizales adyacentes en un bosque lluvioso montano del sur de Costa Rica

Abstract:Successful forest restoration in tropical environments is limited by the paucity of studies on the initial establishment and early survival requirements of seedlings of most native tropical tree species under disturbed conditions. Here, we evaluated the initial growth responses and the regeneration potential of seedlings of five tree species native to Costa Rica (Hasseltia floribunda, Inga densiflora, Persea americana, Tapirira mexicana and Trichilia pittieri). Seedlings were grown in secondary forests and adjacent open pastures under contrasting conditions of light availability. We quantified seedling growth, survival and herbivory from August 2010 to August 2011 on a monthly basis, and measured differences in leaf mass per area (LMA) at the end of the experiment. We found significant variation in growth responses between the understory of secondary forests and pastures. Growth in height was highest in pastures across all species, with I. densiflora, P. americana and T. mexicana showing the most striking differences. In contrast, H. floribunda and T. pittieri did not show differences in diameter growth between environments. Except for T. mexicana, herbivory increased throughout the experiment in all the species. Herbivory increased faster in pastures for H. floribunda and T. pittieri and showed higher rates in the forest understory for I. densiflora and P. americana. Seedling survival showed significant differences for all species across environments. Survival of H. floribunda and I. densiflora was higher in secondary forests, whereas the other species showed higher survival in pastures. LMA showed higher values in the pastures across all species, showing that individuals exposed to high light conditions had great photosynthetic rate and great leaf construction capacity. Due to their rapid growth and high survival, I. densiflora and T. mexicana showed great potential to restore abandoned pastures and secondary forests. Increasing our knowledge on the response of seedlings under disturbed conditions in tropical ecosystems is critical for improving the restoration of altered environments by matching the ecological amplitude of native species with specific environmental conditions. Rev. Biol. Trop. 64 (4): 1565-1583. Epub 2016 December 01.

Resumen:La restauración exitosa del bosque en ambientes tropicales está limitada por la carencia de estudios sobre los requisitos de hábitat, el establecimiento inicial, y la sobrevivencia de las plántulas de la mayoría de las especies de árboles tropicales nativos. En este estudio, evaluamos las respuestas en el crecimiento inicial y el potencial de regeneración de las plántulas de cinco especies de árboles nativos de Costa Rica (Hasseltia floribunda, Inga densiflora, Persea americana, Tapirira mexicana y Trichilia pittieri). Sembramos las plántulas en bosques secundarios y en pastizales abiertos adyacentes con condiciones de luz contrastantes. Cuantificamos el crecimiento, la supervivencia y el daño por herbivoría de las plántulas de forma mensual entre agosto 2010 y agosto 2011, y medimos las diferencias en el la masa foliar por área de la hoja al final del experimento. Existió una variación significativa en el crecimiento de las plántulas expuestas a la sombra en los bosques secundarios y aquellas que crecieron en pastizales. El crecimiento en altura fue mayor en pastizales en todas las especies; I. densiflora, P. americana y T. mexicana mostraron la mayor diferencia entre ambientes. En con- traste, H. floribunda y T. pittieri no mostraron diferencias en el crecimiento del diámetro entre ambientes. Con la excepción de T. mexicana, todas las especies mostraron un aumento en herbivoría durante el estudio. En los pastizales, el incremento de la herbivoría fue más rápido en H. floribunda y T. pittieri, mientras que para I. densiflora y P. americana fue mayor en el bosque. La sobrevivencia de plántulas fue diferente entre ambientes para todas las especies. La sobrevivencia de H. floribunda e I. densiflora fue mayor en el bosque secundario, mientras que en las demás especies hubo mayor sobrevivencia de plántulas en los pastizales. La masa foliar por área fue mayor en las plántulas que crecieron en los pastizales que en bosque secundario para todas las especies, lo que sugiere que los individuos que crecieron en condiciones de alta incidencia de luz tuvieron mayor tasa fotosintética y mayor capacidad de construcción de la hoja. Debido al rápido crecimiento y alta supervivencia I. densiflora y T. mexicana tienen gran potencial para la restauración de bosques secundarios y pastizales abandonados. Generar información sobre la respuesta de plántulas creciendo en bosques secundarios y pastizales en regeneración es crítico para mejorar los procesos de restauración de ambientes alterados. La restauración de estos bosques depende de la correspondencia entre las condiciones específicas de hábitat y la plasticidad ecológica de las especies nativas.

Seedling growth and survival of five tree species in secondary forests and adjacent pastures in the montane rain forests of Southern Costa Rica.

Successful forest restoration in tropical environments is limited by the paucity of studies on the initial establishment and early survival requirements of seedlings of most native tropical tree species under disturbed conditions. Here, we evaluated the initial growth responses and the regeneration potential of seedlings of five tree species native to Costa Rica (Hasseltia floribunda, Inga densiflora, Persea americana, Tapirira mexicana and Trichilia pittieri). Seedlings were grown in secondary forests and adjacent open pastures under contrasting conditions of light availability. We quantified seedling growth, survival and herbivory from August 2010 to August 2011 on a monthly basis, and measured differences in leaf mass per area (LMA) at the end of the experiment. We found significant variation in growth responses between the understory of secondary forests and pastures. Growth in height was highest in pastures across all species, with I. densiflora, P. americana and T. mexicana showing the most striking differences. In contrast, H. floribunda and T. pittieri did not show differences in diameter growth between environments. Except for T. mexicana, herbivory increased throughout the experiment in all the species. Herbivory increased faster in pastures for H. floribunda and T. pittieri and showed higher rates in the forest understory for I. densiflora and P. americana. Seedling survival showed significant differences for all species across environments. Survival of H. floribunda and I. densiflora was higher in secondary forests, whereas the other species showed higher survival in pastures. Leaf mass per area showed higher values in the pastures across all species, it suggests that individuals exposed to high light conditions had great photosynthetic rate and great leaf construction capacity. Due to their rapid growth and high survival, I. densiflora and T. mexicana showed great potential to restore abandoned pastures and secondary forests. Increasing our knowledge on the response of seedlings under disturbed conditions in tropical ecosystems is critical for improving the restoration of altered environments by matching the ecological amplitude of native species with specific environmental conditions.