A transcontinental comparison of the diversity and composition of tropical forest understory herb assemblages

Although tropical forests are renowned for their high plant diversity, to date there has been no global quantitative evaluation of the local species richness of terrestrial forest herbs in tropical forests. In this paper, richness and composition of terrestrial herb assemblages is compared in tropical forests of America, Africa and South East Asia. We established 86 non-continuous transects of 445m each. Herb species richness was analysed and compared to six environmental parameters using minimal adequate regression models and simultaneous autoregressive models. At the global scale, we found a close relationship between herb species richness and temperature parameters, with no differences between continents. The subdivision into three main taxonomic groups (ferns, monocots, dicots) showed that each group has distinct relations to environmental factors and differences in richness between continents. Most of the 72 families found have pantropical distributions but 12, 11, and 16 families were significantly over-represented in America, Africa, and Asia, respectively. Although total species richness was closely related to climatic factors, ferns, monocots and dicots were represented by distinct sets of families with varying species richness on each continent. Which species are found at a given site may thus reflect group-specific evolutionary and historical factor

First record of bat-pollination in the species-rich genus Tillandsia (Bromeliaceae)

Background and Aims Bromeliaceae is a species-rich neotropical plant family that uses a variety of pollinators, principally vertebrates. Tillandsia is the most diverse genus, and includes more than one-third of all bromeliad species. Within this genus, the majority of species rely on diurnal pollination by hummingbirds; however, the flowers of some Tillandsia species show some characteristics typical for pollination by nocturnal animals, particularly bats and moths. In this study an examination is made of the floral and reproductive biology of the epiphytic bromeliad Tillandsia macropetala in a fragment of humid montane forest in central Veracruz, Mexico. Methods The reproductive system of the species, duration of anthesis, production of nectar and floral scent, as well as diurnal and nocturnal floral visitors and their effectiveness in pollination were determined. Key Results Tillandsia macropetala is a self-compatible species that achieves a higher fruit production through outcrossing. Nectar production is restricted to the night, and only nocturnal visits result in the development of fruits. The most frequent visitor (75 % of visits) and the only pollinator of this bromeliad (in 96 % of visits) was the nectarivorous bat Anoura geoffroyi (Phyllostomidae: Glossophaginae). Conclusions This is the first report of chiropterophily within the genus Tillandsia. The results on the pollination biology of this bromeliad suggest an ongoing evolutionary switch from pollination by birds or moths to bat

Biological variation in climbing aroids

Antecedentes y Objetivos: Las especies de Araceae son conocidas por sus distintas formas de vida; sin embargo, la terminología relacionada con este grupo genera confusiones conceptuales dentro de la comunidad científica. Nuestro objetivo fue demostrar la confusión conceptual actual derivada de la literatura y ejemplares de herbario vs. observaciones y datos de campo. Métodos: Se realizó una revisión exhaustiva de fuentes de información relevantes sobre las formas de vida de aráceas, incluyendo floras y tratamientos taxonómicos, así como de ejemplares de herbario. Adicionalmente, se llevó a cabo un censo de seis especies de aráceas en la Estación de Biología Tropical “Los Tuxtlas”, Veracruz, México. Para cada individuo se buscaron las conexiones de sus raíces adventicias y tallos con el suelo, además se tomaron medidas de su longitud total y hojas para crear tres clases ontogénicas. Resultados clave: Se presentan las descripciones de formas de vida de 30 especies de Araceae trepadoras de Veracruz publicadas en nueve publicaciones, mostrando una marcada inconsistencia en el uso de la terminología. El análisis de los datos documentados en los ejemplares de herbario reveló para 17 especies, que todas fueron asignadas con al menos tres de ocho diferentes categorías de hábito, principalmente como epífita, hemiepífita y trepadora. De las seis especies censadas en campo, la mayoría de los individuos tiene conexión con el suelo. Conclusiones: Las asignaciones de formas de vida basadas en observaciones de campo sin tener datos cuantitativos sobre el contacto de las plantas con el suelo son problemáticas. Más bien, es necesario analizar y aclarar la ontogenia de las aráceas trepadoras, lo que puede contribuir a estructurar adecuadamente nuestros conceptos y evitar ambigüedades en la terminología. Asimismo, también se debería evitar que se agrupen a las especies de Araceae indistintamente en estudios sobre epífitas, lo cual lleva a sobreestimar la riqueza reportada.Background and Aims: The species of Araceae are known for their diversity of life forms; however, the terminology regarding this group generates conceptual confusion within the scientific community. Our objective was to demonstrate the current conceptual confusion based on the literature, herbarium vouchers vs. observations and field data. Methods: A thorough review of relevant sources of information on aroid life forms was carried out, including floras and taxonomic treatments, as well as herbarium vouchers. Additionally, a census of six aroid species was conducted at the Tropical Biological Station “Los Tuxtlas”, Veracruz, Mexico. For each individual we searched for the connections of its adventitious roots and stems with the soil, in addition we took measurements of its total length and leaves to create three ontogenic classes. Key results: We present descriptions of life forms of 30 species of climbing Araceae from Veracruz published in nine publications, showing pronounced inconsistency in the use of terminology. The analysis of the data documented in the herbarium specimens revealed for 17 species, that these were assigned to at least three of eight different habit categories, mainly as epiphyte, hemiepiphyte and climber. Of the six species surveyed in the field, most of the individuals have a connection with the soil. Conclusions: Life form assignments based on field observations without quantitative data on the contact of plants with the soil are problematic. Rather, it is necessary to analyze and clarify the ontogeny of climbing aroids, which can help to properly structure our concepts and avoid ambiguities in terminology. Likewise, it should also be avoided that Araceae species are lumped indiscriminately in epiphyte studies, which leads to an overestimation of the reported richness

A transcontinental comparison of the diversity and composition of tropical forest understory herb assemblages

Although tropical forests are renowned for their high plant diversity, to date there has been no global quantitative evaluation of the local species richness of terrestrial forest herbs in tropical forests. In this paper, richness and composition of terrestrial herb assemblages is compared in tropical forests of America, Africa and South East Asia. We established 86 non-continuous transects of 445 m each. Herb species richness was analysed and compared to six environmental parameters using minimal adequate regression models and simultaneous autoregressive models. At the global scale, we found a close relationship between herb species richness and temperature parameters, with no differences between continents. The subdivision into three main taxonomic groups (ferns, monocots, dicots) showed that each group has distinct relations to environmental factors and differences in richness between continents. Most of the 72 families found have pantropical distributions but 12, 11, and 16 families were significantly over-represented in America, Africa, and Asia, respectively. Although total species richness was closely related to climatic factors, ferns, monocots and dicots were represented by distinct sets of families with varying species richness on each continent. Which species are found at a given site may thus reflect group-specific evolutionary and historical factors

Diversidad de arbustos a lo largo de gradientes de elevación y perturbación en el centro de Veracruz, México

Background and Aims: Shrubs are an important floristic element of tropical forests; however, research on plants with this growth form is still scarce. The main objective of this study was to analyze the influence of elevation and disturbance on shrub richness and distribution patterns in the central region of Veracruz, Mexico.Methods: An elevational gradient (20-3500 m a.s.l.) was established along the eastern slopes of the Cofre de Perote, where floristic sampling was conducted at eight study sites within elevational belts of approximately 500 m each. We recorded the abundance of shrubs within 120 plots of 20 × 20 m, distributed in three habitats: conserved, disturbed and secondary forest. The species richness and floristic composition were analyzed, and the resulting data were compared between the different elevational belts and degrees of forest disturbance.Key results: We recorded 118 species and 14 morphospecies of shrubs, representing 8.6% of the shrubs reported for Veracruz. The forest fragments at elevations of 500 and 1000 m are the most species-rich; however, these are the most susceptible to human activities. The anthropogenic disturbance of these forests can increase species richness locally, but at the same time modifies the floristic composition from a conserved habitat to a secondary one.Conclusions: The present study is the first in the Neotropics that considers and analyzes shrubs as an individual group along a complete gradient using a uniform and comparable sampling method. This enabled us to detect which fragments of conserved, disturbed or secondary forest are highly diverse and at the same time susceptible to human disturbance. These sites can serve as strategic buffer zones and refuge of this group of plants, since regardless of their use intensity, they could and should be considered in future conservation plans in this region threatened by environmental degradation.Antecedentes y Objetivos: Los arbustos son un elemento florístico importante de los bosques tropicales; sin embargo, la investigación sobre plantas con esta forma de crecimiento aún es escasa. El objetivo principal de este estudio consistió en analizar la influencia de la elevación y la perturbación en los patrones de distribución y diversidad de los arbustos en la región central de Veracruz, México.Métodos: Se estableció un gradiente elevacional (20-3500 m s.n.m) en la ladera este del Cofre de Perote, donde se realizó un muestreo florístico en ocho pisos elevacionales separados por 500 m cada uno. Registramos la abundancia de arbustos dentro de 120 parcelas de 20 × 20 m, distribuidas en tres hábitats: bosque conservado, perturbado y secundario. Se analizó la riqueza, diversidad y composición florística; además, se compararon los datos resultantes entre los diferentes pisos elevacionales y los grados de perturbación del bosque.Resultados clave: Se registraron 118 especies y 14 morfoespecies de arbustos, lo que representa 8.6% de los reportados para Veracruz. Los fragmentos de bosque en las elevaciones de 500 y 1000 m son los más ricos en número de especies, pero también son los más susceptibles a las perturbaciones humanas. La influencia antrópica sobre estos bosques puede incrementar la diversidad de especies a nivel local, pero al mismo tiempo modifica la composición florística de un hábitat conservado a uno secundario.Conclusiones: El presente estudio es el primero en el Neotrópico en considerar y analizar los arbustos como un grupo individual a lo largo de un gradiente completo utilizando un método de muestreo uniforme y comparable. Esto permitió detectar aquellos fragmentos de bosque conservado, perturbado o secundario que son altamente diversos y susceptibles a los distrubios antrópicos. Estos sitios sirven como zonas estratégicas de amortiguación y refugio de este grupo de plantas, ya que independientemente de la intensidad de su uso, estos podrían y deberían considerarse en futuros planes de conservación en esta región amenazada por la degradación ambiental

New Proposal of Epiphytic Bromeliaceae Functional Groups to Include Nebulophytes and Shallow Tanks

The Bromeliaceae family has been used as a model to study adaptive radiation due to its terrestrial, epilithic, and epiphytic habits with wide morpho-physiological variation. Functional groups described by Pittendrigh in 1948 have been an integral part of ecophysiological studies. In the current study, we revisited the functional groups of epiphytic bromeliads using a 204 species trait database sampled throughout the Americas. Our objective was to define epiphytic functional groups within bromeliads based on unsupervised classification, including species from the dry to the wet end of the Neotropics. We performed a hierarchical cluster analysis with 16 functional traits and a discriminant analysis, to test for the separation between these groups. Herbarium records were used to map species distributions and to analyze the climate and ecosystems inhabited. The clustering supported five groups, C3 tank and CAM tank bromeliads with deep tanks, while the atmospheric group (according to Pittendrigh) was divided into nebulophytes, bromeliads with shallow tanks, and bromeliads with pseudobulbs. The two former groups showed distinct traits related to resource (water) acquisition, such as fog (nebulophytes) and dew (shallow tanks). We discuss how the functional traits relate to the ecosystems inhabited and the relevance of acknowledging the new functional groups

Phylogenetic diversity and the structure of host-epiphyte interactions across the Neotropics

Understanding the mechanisms driving community assembly has been a major focus of ecological research for nearly a century, yet little is known about these mechanisms in commensal communities, particularly with respect to their historical/evolutionary components. Here, we use a large-scale dataset of 4,440 vascular plant species to explore the relationship between the evolutionary distinctiveness (ED) (as measured by the 'species evolutionary history' (SEH)) of host species and the phylogenetic diversity (PD) of their associated epiphyte species. Although there was considerable variation across hosts and their associated epiphyte species, they were largely unrelated to host SEH. Our results mostly support the idea that the determinants of epiphyte colonization success might involve host characteristics that are unrelated to host SEH (e.g., architectural differences between hosts). While determinants of PD of epiphyte assemblages are poorly known, they do not appear to be related to the evolutionary history of host species. Instead, they might be better explained by neutral processes of colonization and extinction. However, the high level of phylogenetic signal in epiphyte PD (independent of SEH) suggests it might still be influenced by yet unrecognized evolutionary determinants. This study highlights how little is still known about the phylogenetic determinants of epiphyte communities

EpIG‐DB: A database of vascular epiphyte assemblages in the Neotropics

Vascular epiphytes are a diverse and conspicuous component of biodiversity in tropical and subtropical forests. Yet, the patterns and drivers of epiphyte assemblages are poorly studied in comparison with soil‐rooted plants. Current knowledge about diversity patterns of epiphytes mainly stems from local studies or floristic inventories, but this information has not yet been integrated to allow a better understanding of large‐scale distribution patterns. EpIG‐DB, the first database on epiphyte assemblages at the continental scale, resulted from an exhaustive compilation of published and unpublished inventory data from the Neotropics. The current version of EpIG‐DB consists of 463,196 individual epiphytes from 3,005 species, which were collected from a total of 18,148 relevés (host trees and ‘understory’ plots). EpIG‐DB reports the occurrence of ‘true’ epiphytes, hemiepiphytes and nomadic vines, including information on their cover, abundance, frequency and biomass. Most records (97%) correspond to sampled host trees, 76% of them aggregated in forest plots. The data is stored in a TURBOVEG database using the most up‐to‐date checklist of vascular epiphytes. A total of 18 additional fields were created for the standardization of associated data commonly used in epiphyte ecology (e.g. by considering different sampling methods). EpIG‐DB currently covers six major biomes across the whole latitudinal range of epiphytes in the Neotropics but welcomes data globally. This novel database provides, for the first time, unique biodiversity data on epiphytes for the Neotropics and unified guidelines for future collection of epiphyte data. EpIG‐DB will allow exploration of new ways to study the community ecology and biogeography of vascular epiphytes

The role of hummingbirds in the evolution and diversification of Bromeliaceae: unsupported claims and untested hypotheses

At least half of the 3600 species of Bromeliaceae are pollinated by hummingbirds. There is little doubt that the four to 12 evolutionary shifts towards and c. 32 shifts away from hummingbird pollination opened new evolutionary spaces for bromeliad diversification, and that hummingbird pollination has led to increased bromeliad diversification rates. However, the mechanisms leading to these increased rates remain unclear. We here propose that there are four main types of mechanisms that may increase diversification rates of hummingbird-pollinated bromeliad clades: (1) bromeliad speciation through adaptation to different hummingbird species; (2) increased allopatric speciation in hummingbird-pollinated clades due to lower pollen transfer efficiency compared with other pollinators; (3) differential speciation rates in hummingbird-pollinated clades dependent on of flowering phenology and hummingbird behaviour; and (4) higher speciation rates of bromeliads in montane environments (where hummingbird pollination predominates) due to topographic population fragmentation. To date, none of these hypotheses has been appropriately tested, partly due to a lack of data, but also because research so far has focused on documenting the pattern of increased diversification in hummingbird-pollinated clades, implicitly assuming that this pattern supports an underlying mechanism while ignoring the fact that several competing mechanisms may be considered. The aim of the present review is to increase awareness of these mechanisms and to trigger research aimed at specifically testing them. We conclude that much additional research on the roles of hummingbird behaviour and gene flow between bromeliad species is needed to elucidate their contribution to the evolution of diversity in bromeliads and other plant families