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This study characterizes the osmophores and corolla traits in 18 species of Bignonieae Dumort., a Bignoniaceae tribe occurring in the Cerrado, a neotropical savanna in Brazil. To detect osmophore distribution, whole, newly opened flowers were immersed in Neutral Red Solution. Samples from the corolla tube and lobes were also fixed and analyzed micromorphologically, anatomically, and histochemically. The osmophores showed six markedly different distribution patterns that were not clearly associated with histological features. In most species, osmophores comprised papillose secretory epidermises and a few layers of subepidermal parenchyma. Starch grains, lipid droplets, and terpenes were detected in osmophores. An ornamented cuticle, cuticular folds, glandular and non-glandular trichomes, raised stomata and epicuticular wax granules are common traits in the species studied and may be useful in determining the taxonomy of the group. We found that 94% of the species visited by bees had papillose epidermises while the single hummingbird-pollinated species presented a flattened epidermis. Variations in osmophore pattern among species visited by bees, including variations within the same plant genus, are novel finding. Additionally, the Bignonieae species visited by bees presented a textured corolla surface, which has been reported as facilitating bee attachment and movement towards the floral resource. Future studies with a greater number of Bignonieae species and more detailed pollinator behavioral assays may help in the interpretation of the variations in corolla traits and functional relationships between flowers and pollinators.

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We have previously suggested that a shift from bee to hummingbird pollination, in concert with floral architecture modifications, occurred at the crown of Salvia subgenus Calosphace in North America ca. 20 mya (Kriebel et al. 2020 and references therein). Sazatornil et al. (2022), using a hidden states model, challenged these assertions, arguing that bees were the ancestral pollinator of subg. Calosphace and claiming that hummingbirds could not have been the ancestral pollinator of subg. Calosphace because hummingbirds were not contemporaneous with crown subg. Calosphace in North America. Here, using a variety of models, we demonstrate that most analyses support hummingbirds as ancestral pollinators of subg. Calosphace and show that Sazatornil et al. (2022) erroneously concluded that hummingbirds were absent from North America ca. 20 mya. We contend that "biological realism" - based on timing and placement of hummingbirds in Mexico ca. 20 mya and the correlative evolution of hummingbird associated floral traits - must be considered when comparing models based on fit and complexity, including hidden states models.

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Many flowers are fed on by florivores, but we know little about if and how feeding on flowers affects their visual and chemical advertisement and nectar resource, which could disrupt pollination. Here, we investigated if damages caused by florivores compromise a Neotropical hummingbird pollination system, by modifying the floral advertisements and the nectar resource. We surveyed natural florivory levels and patterns, examined short-term local effects of floral damages caused by the most common florivore, a caterpillar, on floral outline, intra-floral colour pattern and floral scent, as well as on the amount of nectar. Following, we experimentally tested if the most severe florivory pattern affected hummingbird pollination. The feeding activity of the most common florivore did not alter the intra-floral colour pattern, floral scent, and nectar volume, but changed the corolla outline. However, this change did not affect hummingbird pollination. Despite visual floral cues being important for foraging in hummingbirds, our results emphasise that changes in the corolla outline had a neutral effect on pollination, allowing the maintenance of florivore-plant-pollinator systems without detriment to any partner.

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Rising temperatures and increasing drought in Mediterranean-type climate areas are expected to affect plant-pollinator interactions, especially in plant species with specialised pollination. Central Chile experienced a mega drought between 2010 and 2020 which reached an extreme in the austral summer of 2019-2020. Based on intensive pollinator sampling and floral studies we show that the subalpine form of Mutisia subulata (Asteraceae) is a specialised hummingbird-pollinated species. In a two-year study which included the severest drought year, we quantified visitation frequency, flower-head density, flower-head visitation rates, two measures of floral longevity, nectar characteristics and seed set and monitored climatic variables to detect direct and indirect climate-related effects on pollinator visitation. Flower-head density, nectar standing crop and seed set were significantly reduced in the severest drought year while nectar concentration increased. The best model to explain visitation frequency included flower-head density, relative humidity, temperature, and nectar standing crop with highly significant effects of the first three variables. Results for flower-head density suggest hummingbirds were able to associate visual signals with reduced resource availability and/or were less abundant. The negative effect of lower relative humidity suggests the birds were able to perceive differences in nectar concentration. Reduced seed set per flower-head together with the availability of far fewer ovules in the 2019-2020 austral summer would have resulted in a major reduction in seed set. Longer and more intense droughts in this century could threaten local population persistence in M. subulata.

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PREMISE: Closely related plant species with overlapping ranges often experience competition for pollination services. Such competition can select for divergence in floral traits that attract pollinators or determine pollen placement. While most species in Centropogon (Campanulaceae: Lobelioideae) have flowers that suggest adaptation to bat or hummingbird pollination, actual pollinators are rarely documented, and a few species have a mix of traits from both pollination syndromes. We studied the pollination biology of a "mixed-syndrome" species and its co-occurring congeners to examine the relationship between floral traits and visitation patterns for Centropogon. METHODS: Fieldwork at two sites in Bolivian cloud forests involved filming floral visitors, quantifying pollen transfer, and measuring floral traits. Stamen exsertion, which determines pollen placement, was measured from herbarium specimens across the geographic range of these species to test for character displacement. RESULTS: Results show a generalization gradient, from primarily bat pollination in white-flowered Centropogon incanus, to bat pollination with secondary hummingbird pollination in the cream-flowered C. brittonianus, to equal reliance on both pollinators in the red-flowered, mixed-syndrome C. mandonis. Pollen transfer between these species is further reduced by differences in stamen exsertion that are accentuated in zones of sympatry, a pattern consistent with character displacement. CONCLUSIONS: Our results demonstrate that key differences in floral color and shape mediate a gradient of specialization in Bolivian Centropogon. Interspecific pollen transfer is further reduced by potential character displacement of a key trait. Broadly, our results have implications for understanding the hyper-diversity of Andean cloud forests, in which multiple species of the same genus frequently co-occur.

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In tropical trees, forest fragmentation has been shown to affect mating and gene flow patterns. Mobile dispersal vectors should be less sensitive to fragmented landscapes and may ameliorate the genetic effects of forest fragmentation on plant populations. To test this hypothesis, we analyzed gene flow patterns in Symphonia globulifera, a tropical tree species with highly mobile pollinators and seed dispersers in the Osa Peninsula in southern Costa Rica. We used microsatellites to study genetic diversity and realized gene flow patterns between a continuous forest and a forest fragment. We found high levels of genetic diversity in adults and seedlings at both sites. Parentage analyses suggest near-neighbor matings and frequent long-distance gene flow events. Half the progeny beneath an adult was not sired by that tree and the majority of established seedlings were the result of long-distance gene dispersal. Gene flow from the forest into the fragment was more common than from the fragment into the continuous forests. Despite long distance gene flow events, seedling spatial genetic structure was stronger and extended further in the forest fragment likely due to limited seed dispersal. We conclude that fragmentation affects gene flow in this tropical tree and may compromise its genetic diversity in forest fragments even for a species with mobile pollen and seed vectors.

En árboles tropicales se ha demostrado que la fragmentación del bosque afecta los patrones de apareamiento y reduce el flujo génico. Las especies forestales cuyos dispersores y polinizadores son capaces de moverse largas distancias sobre ambientes alterados, serán menos sensibles a los efectos de la fragmentación y podrán intercambiar genes regularmente entre fragmentos. Para probar esta hipótesis, en la Península de Osa en el sur de Costa Rica, estudiamos los patrones de flujo génico de Symphonia globulifera, una especie de árbol tropical con polinizadores y dispersores de semillas altamente móviles. Utilizamos microsatélites para estudiar la diversidad genética y los patrones de flujo génico entre un bosque continuo y un fragmento de bosque. Encontramos altos niveles de diversidad genética tanto en adultos como en plántulas. Los análisis de parentesco indican apareamiento entre vecinos cercanos, sin embargo el flujo génico a larga distancia es frecuente. La mitad de la progenie debajo de un adulto no es engendrada por ese árbol y la mayoría de las plántulas establecidas son el resultado del flujo génico a larga distancia. El flujo de genes desde el bosque continuo hacia el fragmento era más frecuente que el movimiento inverso. A pesar del flujo génico a larga distancia, hay diferencias en frecuencias alélicas entre plántulas y la estructura espacial era más fuerte y se extendía a mayor distancia en el fragmento, probablemente por una dispersión limitada de semillas. Concluimos que la fragmentación afecta los patrones de flujo génico en esta especie de árbol tropical y puede afectar su diversidad genética en paisajes fragmentados, inclusive para una especies con polinizadores y dispersores con alta movilidad.

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AbstractPitcairnioideae is the second most diverse subfamily of bromeliads (Bromeliaceae), a group exclusive to tropical regions of the New World. Pitcairnioid bromeliads have floral traits assumed to promote outcrossing through biotic pollination systems; however, the reproductive biology of most of the species of this group has not been documented. Pitcairnia heterophylla is an epiphytic (seldom saxicolous) bromeliad occurring from Southern Mexico, into the Northern Andes. We studied the pollination and breeding system of P. heterophylla in an epiphytic population at a mountain forest in Costa Rica from January to April 2013. We performed hand pollination experiments (agamospermy, autonomous self-pollination, hand self-pollination and hand cross-pollination) on 89 flowers from 23 individuals (3-6 flowers per individual) in 2013 flowering season. Nectar production was measured on 18 unvisited flowers of six individuals with a hand-held refractometer. Simultaneously, floral visitors were recorded on eight individuals with trail cameras for a total of 918 hours (115 ± 52 hours per individual, mean ± SE). Under natural conditions, seed set (540.4 ± 55.2) was similar to manually selfed flowers (516.3 ± 41.5) and autonomously selfed flowers (521.1 ± 29.0), but lower to manually outcrossed flowers (670.2 ± 31.3). The flowers of P. heterophylla are self-compatible, capable of autonomous pollination, and non-agamospermous. Intrafloral self-pollination is facilitated by adichogamy and lack of floral herkogamy. The scentless red flowers of P. heterophylla with tubular corollas and nectar production suggested ornithophilic pollination which was confirmed by video recording of 46 hummingbird visits. The most common floral visitor was the short-billed hummingbird Lampornis calolaemus which accounted for 78 % of the visits. However, the visitation rate during the flowering season was low (0.6 visits per day per plant). Selfing in P. heterophylla might be explained as a mechanism of reproductive assurance and to reduce interspecific pollen flow with taxonomically unrelated plants.

ResumenLa subfamilia Pitcairnioideae es la segunda más diversa de las bromelias (Bromeliaceae), un grupo exclusivo de la región Neotropical. Las bromelias pitcairnioideas poseen rasgos florales que se asume promueven el exo-cruzamiento a través de sistemas de polinización biótica; sin embargo, la biología reproductiva de la mayoría de especies de este grupo no se ha documentado. Pitcairnia heterophylla es una bromelia epífita (raramente saxícola) que se encuentra en el sur de México, América Central y el norte de los Andes. Se estudió la polinización y el sistema de apareamiento de una población P. heterophylla en un bosque montano en Costa Rica entre enero y abril 2013. Se realizaron cuatro experimentos de polinización (agamospermia, auto-polinización espontánea, auto-polinización manual, exo-polinización manual) en 89 flores de 23 individuos (3-6 flores por individuo). Se cuantificó la producción de néctar de 18 flores en seis individuos con un refractómetro. Simultáneamente, se registraron los visitantes florales de ocho individuos en el campo con ayuda de cámaras trampa por un total de 918 horas (115 ± 52 horas por individuo, promedio ± ES). Bajo condiciones naturales la producción de semillas fue similar (540.4 ± 55.2) a la producción de las flores auto-polinizadas manualmente (516.3 ± 41.5) y las flores auto-polinizadas espontáneamente (521.1 ± 29.0), pero fue menor a las flores exo-cruzadas (670.2 ± 31.3). Las flores de P. heterophylla son auto-compatibles, con alta capacidad de polinización espontánea y sin capacidad de agamospermia. La auto-polinización intra-floral es facilitada por la ausencia de dicogamia y hercogamia floral. Las flores rojas, sin aroma, con corola tubular y con producción de néctar sugirieron polinización por ornitofilia, lo que se confirmó con el registro de 46 visitas por colibríes (Apodiformes: Trochilidae). El visitante floral más común fue Lampornis calolaemus (78 % de las visitas). A pesar de esto, la tasa de visitas durante el periodo de estudio fue baja (0.6 visitas por día por planta). La capacidad de autofecundación de P. heterophylla puede ser explicada como un mecanismo de aseguramiento reproductivo, así como para reducir el flujo de polen inter-específico de plantas taxonómicamente no relacionadas.

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Many mistletoe species produce 'bird'-pollinated flowers; however, the reproductive biology of the majority of these species has not been studied. Psittacanthus auriculatus is a Mexican endemic mistletoe, most common in open, dry mesquite grassland. Knowledge of the reproductive biology of P. auriculatus is essential for understanding species formation and diversification of Psittacanthus mistletoes, but it is currently poorly understood. Thus, we studied floral biology and phenology, nectar production and breeding system and pollination of this species. The hermaphroditic red-pink flowers open from the middle to the tip and petals are curly, but remain partially fused forming a floral tube of ca. 20-mm long. Flowers are partially protandrous, produce large amounts of nectar, last 2 days, and stigma receptivity is highest during the second day. We recorded hummingbirds (Cynanthus latirostris, Hylocharis leucotis, Amazilia beryllina, A. violiceps, Calothorax lucifer, Archilochus colubris) and less commonly butterflies (Agraulis vanillae, Anteos clorinde, Papilio multicaudatus, Phocides urania, Phoebis sennae) as floral visitors. P. auriculatus flowers are self-compatible. However, this mistletoe is an obligate animal-pollinated species, as the sensitive stigma avoids self-pollination. Under natural conditions, reproductive success was higher than in manually selfed or cross-pollinated flowers, likely due to the traplining foraging behaviour of hummingbirds. We suggest that the apparent efficient foraging behaviour of hummingbirds maintains gene flow among P. auriculatus, promoting outcrossing.

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