Tilo Henning

Diversity patterns of selected Andean plant groups correspond to topography and habitat dynamics, not orogeny

The tropical Andes are a hotspot of biodiversity, but detailed altitudinal and latitudinal distribution patterns of species are poorly understood. We compare the distribution and diversity patterns of four Andean plant groups on the basis of georeferenced specimen data: the genus Nasa (Loasaceae), the two South American sections of Ribes (sect. Parilla and sect. Andina, Grossulariaceae), and the American clade of Urtica (Urticaceae). In the tropical Andes, these often grow together, especially in (naturally or anthropogenically) disturbed or secondary vegetation at middle to upper elevations. The climatic niches of the tropical groups studied here are relatively similar in temperature and temperature seasonality, but do differ in moisture seasonality. The Amotape–Huancabamba Zone (AHZ) between 3 and 8° S shows a clear diversity peak of overall species richness as well as for narrowly endemic species across the groups studied. For Nasa, we also show a particular diversity of growth forms in the AHZ. This can be interpreted as proxy for a high diversity of ecological niches based on high spatial habitat heterogeneity in this zone. Latitudinal ranges are generally larger toward the margins of overall range of the group. Species number and number of endemic species of our taxa peak at elevations of 2,500–3,500 m in the tropical Andes. Altitudinal diversity patterns correspond well with the altitudinal distribution of slope inclination. We hypothesize that the likelihood and frequency of landslides at steeper slopes translate into temporal habitat heterogeneity. The frequency of landslides may be causally connected to diversification especially for the numerous early colonizing taxa, such as Urtica and annual species of Nasa. In contrast to earlier hypotheses, uplift history is not reflected in the pattern here retrieved, since the AHZ is the area of the most recent Andean uplift. Similarly, a barrier effect of the low-lying Huancabamba depression is not retrieved in our data.

Total control – pollen presentation and floral longevity in Loasaceae (Blazing Star Family) are modulated by light, temperature and pollinator visitation rates

Stamen movements can be understood as a mechanism influencing pollen presentation and increasing outbreeding success of hermaphroditic flowers via optimized male function. In this study we experimentally analyzed the factors regulating autonomous and thigmonastic (triggered by flower visitors) stamen movements in eight species of Loasaceae. Both types of stamen movements are positively influenced by light and temperature and come to a virtual standstill in the dark and at low temperatures (12°C). Pollen presentation is thus discontinued during periods where pollinators are not active. Overall stamen presentation increases with increasing flower age. Contrary to expectation, no geometrical correlation between the floral scale stimulated and the stamen fascicle reacting exists, indicating that the stimulus is transmitted over the receptacle and stamen maturation dictates which and how many stamens react. Thigmonastic stamen presentation is dramatically accelerated compared to autonomous movement (3–37 times), indicating that the rate of stamen maturation can be adjusted to different visitation schedules. Flowers can react relatively uniformly down to stimulation intervals of 10–15 min., consistently presenting comparable numbers of stamens in the flower c. 5 min. after the stimulus and can thus keep the amount of pollen presented relatively constant even under very high visitation frequencies of 4–6 visits/h. Thigmonastic pollen presentation dramatically reduces the overall duration of the staminate phase (to 1/3rd in Nasa macrothyrsa). Similarly, the carpellate phase is dramatically reduced after pollination, down to 1 d from 4 d. Overall flower longevity is reduced by more than 2/3rds under high visitation rates (<3 d versus 10 d under visitor exclusion) and depleted and pollinated flowers are rapidly removed from the pool. Complex floral behaviour in Loasaceae thus permits a near-total control over pollen dispensation schedules and floral longevity of the individual flower by an extraordinary fine-tuning to both biotic and abiotic factors.

Sample data processing in an additive and reproducible taxonomic workflow by using character data persistently linked to preserved individual specimens

We present the model and implementation of a workflow that blazes a trail in systematic biology for the re-usability of character data (data on any kind of characters of pheno- and genotypes of organisms) and their additivity from specimen to taxon level. We take into account that any taxon characterization is based on a limited set of sampled individuals and characters, and that consequently any new individual and any new character may affect the recognition of biological entities and/or the subsequent delimitation and characterization of a taxon. Taxon concepts thus frequently change during the knowledge generation process in systematic biology. Structured character data are therefore not only needed for the knowledge generation process but also for easily adapting characterizations of taxa. We aim to facilitate the construction and reproducibility of taxon characterizations from structured character data of changing sample sets by establishing a stable and unambiguous association between each sampled individual and the data processed from it. Our workflow implementation uses the European Distributed Institute of Taxonomy Platform, a comprehensive taxonomic data management and publication environment to: (i) establish a reproducible connection between sampled individuals and all samples derived from them; (ii) stably link sample-based character data with the metadata of the respective samples; (iii) record and store structured specimen-based character data in formats allowing data exchange; (iv) reversibly assign sample metadata and character datasets to taxa in an editable classification and display them and (v) organize data exchange via standard exchange formats and enable the link between the character datasets and samples in research collections, ensuring high visibility and instant re-usability of the data. The workflow implemented will contribute to organizing the interface between phylogenetic analysis and revisionary taxonomic or monographic work. Database URL: http://campanula.e-taxonomy.net/

Beautiful, complicated—and intelligent? Novel aspects of the thigmonastic stamen movement in Loasaceae

In a recent study we investigated the complex mechanisms regulating the pollen release via thigmonastic stamen movement found exclusively in Loasaceae subfamily Loasoideae. We demonstrated that stamen movement is modulated by abiotic (light and temperature) as well as biotic stimuli (pollinator availability and visitation frequency). This is explained as a mechanism to adjust the rate of stamen movement and thus pollen dispensation to different environmental conditions in order to optimize pollen transfer. Stamen movement is rapid and thus a near-immediate response to pollinator visits. However, Loasaceae flowers also show a response to biotic stimuli on a longer time scale, by adjusting the duration of both the staminate and the carpellate phase of the anthesis. We here present two additional data sets on species not previously studied, underscoring the shortening of the staminate phase in the presence of pollinator visits vs. their absence and the shortening of the carpellate phase after pollination. Overall, the plant shows not only a rapid but an “intelligent” reaction to its environment in adjusting anthesis and pollen presentation to a range of factors. The physiological and morphological bases of the stamen movement are poorly understood. Our previous study showed that there is no direct spatial relationship between the place of stimulation in the flower and the stamen bundle activated. We here further show the morphological basis for stamen movement from a reflexed into an erect position: Only the basal part of the filament curves around the receptacle, while the upper part of the filament retains its shape. We hypothesize that the stimulus is transmitted over the entire receptacle and the place of reaction is determined by stamen maturity, not the location of the stimulus.

A revision of Nasa ser. Carunculatae (Loasaceae subfam. Loasoideae)

Abstract Nasa Weigend ser. Carunculatae (Urb. & Gilg) Weigend is revised, four species are recognized, and one, Nasa usquiliensis Weigend, T. Henning & C. Schneider described as new to science. The species of this group are restricted to steep scree slopes in Peru and southern Ecuador and have patchy distributions at elevations of 2200–3500 m. All species share shrubby habit with ligneous stems and deciduous foliage. These characters in combination with clawed, spreading petals and bicolored nectar scales are unique in Nasa. Branched root-tubers are reported from N. carunculata, which is the first report of tuberous storage roots in the genus. Species delimitation is primarily based on habit, trichome characters, flower size, petal color and shape, and the morphology of the floral scales.

Dynamic diversification history with rate upshifts in Holarctic bell‐flowers (Campanula and allies)

Campanula s.l. is one of the most speciose flowering plant lineages of the Holarctic (ca. 600 species). In the present study we sequenced three regions of the plastid genome (petD, rpl16 and trnK/matK) across a broad sample of Campanula s.l., which markedly improved phylogenetic resolution and statistical support compared to previous studies. Based on this robust phylogenetic hypothesis we estimated divergence times using BEAST, diversification rate shifts using Bayesian Analysis of Macroevolutionary Mixture (BAMM) and TreePar, and ancestral ranges using Biogeography with Bayesian (and likelihood) Evolutionary Analyses in R. Campanula s.l. is estimated to have originated during the Early Eocene but the major diversification events occurred between the Late Oligocene and Middle Miocene. Two upward diversification rate shifts were revealed by BAMM, specific to the crown nodes of two Campanula clades: CAM17, a mostly South European‐SW Asian lineage originating during the Middle Miocene and containing nearly half of all known Campanula species; and CAM15B, a SW Asian–Sino‐Himalayan lineage of nine species originating in the early Pleistocene. The dynamic diversification history of Campanula and the inferred rate shifts are discussed in a geo‐historical context.

Building compatible and dynamic character matrices – Current and future use of specimen-based character data

Abstract Herbarium specimens have always played a central role in plant sciences and constitute the cornerstone for systematic and taxonomy. This role is further strengthened with the ongoing digitisation and growing online-availability of collections all over the globe. The increasing usability of specimens demands, however, an improved use and sustainable handling of specimen data not only in new scientific uses correlated with the digitisation, but also by modern workflows applied to the traditional purpose of specimens. A crucial step in the comparative analyses of organisms is the preparation of a character matrix to observe and assess the morphological extent and variability of taxa on the basis of individual specimens. This process and the resulting matrix often are of ephemeral nature since only its results are published in a condensed form. The data relationships are usually not stored, making a re-use impossible and a new analysis inevitable. To overcome the limitations of conventional taxonomy, we here introduce a comprehensive workflow that is currently being implemented on the EDIT Platform for Cybertaxonomy.

A case of behavioural diversification in male floral function – the evolution of thigmonastic pollen presentation

Obvious movements of plant organs have fascinated scientists for a long time. They have been studied extensively, but few behavioural studies to date have dealt with them, and hardly anything is known about their evolution. Here, we present a large experimental dataset on the stamen movement patterns found in the Loasaceae subfam. Loasoideae (Cornales). An evolutionary transition from autonomous-only to a combination of autonomous and thigmonastic stamen movement with increased complexity was experimentally demonstrated. We compare the stamen movement patterns with extensive pollinator observations and discuss it in the context of male mating behavior. Thigmonastic pollen presentation via stamen movements appears to be a crucial component of floral adaptation to pollinator behaviour, evolving in concert with complex adjustments of flower signal, reward and morphology. We hypothesize that rapid adjustments of pollen presentation timing may play a significant role in the diversification of this plant group, representing a striking example for the evolutionary significance of plant behaviour.