Bird Pollination Research Articles

Why is pollen in Camellia oleifera inedible to honeybees?

This Commentary examines a recent study that addressed a long-standing controversy: Is the lethal effect of Tea-oil Camellia on honeybee larvae due to nectar or pollen toxicity? Flowers of Camellia oleifera are adapting to bird pollination, evolving 'anti-bee' traits such as theasaponin-containing pollen, which is toxic to bee larvae.

Equivalent mating system parameters in post‐mining and undisturbed native plant populations confirms restitution of bird‐pollinator function

Abstract The restoration of diverse self‐sustaining ecosystems requires re‐establishment of functional interactions among species. For plant communities, pollinators are usually essential for pollination, seed set and seed quality. A common assumption in ecological restoration for plants pollinated by animals is one of ‘build it and they will come’, which is rarely tested. Beyond seed set, there may be negative genetic consequences for seed quality if pollinators and their behaviour do not reflect those in reference populations. Here, we conduct an ecological genetic assessment of seed quality via mating system parameters in Lambertia multiflora (Proteaceae), a species dependent on nectar‐feeding birds for pollination. Four populations of L. multiflora in disturbed sites that were rehabilitated following mineral sands mining were compared with four native reference populations, near Eneabba, Western Australia. In each population, approximately 10 offspring from each of 10 maternal plants were genotyped with 11 highly polymorphic microsatellite markers. From these data, genetic diversity and mating system parameters were assessed, and found to be equivalent across all populations. Mean allelic diversity and heterozygosity across loci were very high. All populations were completely outcrossing with no bi‐parental inbreeding. Mean correlated paternity, sibship and effective population size estimates for restored and natural populations were not significantly different and reflected uniformly high paternal diversity and wide outcrossing. Equivalent genetic results for restored and natural reference populations indicate successful restitution of bird‐pollinator services for L. multiflora in these post‐mining rehabilitation sites. Synthesis and applications. Reviewing our results with other published studies to date suggests a resilience of bird‐pollinator services in restored plant communities. These findings provide some reassurance to restoration practitioners working in these global south systems where bird pollination is a feature, at least for similar landscape scenarios. Our study also highlights the global contribution of ecological genetics to the objective assessment of functional species interactions in ecological restoration, an increasingly important goal of land managers and regulators seeking to improve restoration standards.

Strong habitat and seasonal phenology effects on the evolution of self-compatibility, clonality and pollinator shifts in Lachenalia (Asparagaceae: Scilloideae).

Plants employ a diversity of reproductive safeguarding strategies to circumvent the challenge of pollen limitation. Focusing on southern African Lachenalia (Asparagaceae: Scilloideae), we test the hypothesis that the evolution of reproductive safeguarding traits (self-compatibility, autonomous selfing, bird pollination and clonal propagation) is favoured in species occupying conditions of low insect abundance imposed by critically infertile fynbos heathland vegetation and by flowering outside the austral spring insect abundance peak. We trace the evolution of these traits and selective regimes on a dated, multi-locus phylogeny of Lachenalia and assess their evolutionary associations using ordinary and phylogenetic regression. Ancestral state reconstructions identify an association with non-fynbos vegetation and spring flowering as ancestral in Lachenalia, the transition to fynbos vegetation and non-spring flowering taking place multiple times. They also show that self-compatibility, autofertility, bird pollination and production of multiple clonal offsets have evolved repeatedly. Regression models suggest that bird pollination and self-compatibility are selected for in fynbos and in non-spring flowering lineages, with autofertility being positively associated with non-spring flowering. These patterns support the interpretation of these traits as reproductive safeguarding adaptations under reduced insect pollinator abundance. We find no evidence to support the interpretation of clonal propagation as a reproductive safeguarding strategy.

Species diversity and extinction risk of vertebrate pollinators in India

Animal pollinators underpin the functioning and persistence of ecosystems globally. However, the vital role of pollination is being progressively eroded by the worldwide decline of pollinator species caused by human-induced environmental degradation, resulting in rising costs to biodiversity, agriculture, and economy. Most studies quantifying pollinator diversity and declines have focused on insects, whereas vertebrate pollinators remain comparatively neglected. Here, we present the first comprehensive study quantifying the macroecological patterns of species richness and extinction risk of bird and mammal pollinators in India, a region of extremely high biodiversity and increasing anthropogenic pressure. Our results reveal that hotspots of mammal pollinator diversity are restricted to the south of the Western Ghats, whereas bird pollinator diversity hotspots are scattered throughout the country. Analyses of hotspots of threatened species (based on the IUCN Red List) show that only mammal pollinators are currently classified as threatened in India, whereas multiple hotspots of population declines were observed for birds, and primarily in the Southwest for mammal pollinators. Our analyses failed to identify a role for species traits as drivers of these patterns, whereas most pollinators appear to be threatened by agriculture, logging and hunting for food, and medicinal purposes. Pollinator endangerment has widescale ecological and economic implications such as reduced food production, plant extinction, loss of functional and genetic diversity, and economic damage. We suggest protection of vertebrate pollinators should be emphasised in active conservation agendas in India.

Natural history of pollination of an Asian bignoniaceous tree: The long-tubed flower and the not-so-long bird bill

Bird pollination is well-established in New World Bignoniaceae, but studies of species with floral traits suggestive of bird pollination in the Old World are lacking. Here we studied the pollination ecology of Mayodendron igneum to test the prediction of pollination by specialist flower-visiting birds. Observations from multiple sites showed that both the Streaked Spiderhunter (Arachnothera magna) and pollen-collecting bees were floral visitors. However, almost no fruit was produced if birds were excluded, suggesting that bees do not play a role in pollination, and that pollination is performed almost exclusively by birds in this self-incompatible tree. Measurements of floral traits revealed a typical bird pollination syndrome, and the nectar concentration and volume were both within the proposed ranges based on other flowers pollinated by specialist birds. However, the rather low level of sucrose (less than 2 %) in nectar sugar contradicts the expectation for nectar of flowers pollinated by specialist nectar-feeding birds. Although the Streaked Spiderhunter is among the longest-billed flower-visiting birds in Asia, its bill is only 2/3 of the corolla tube in length, suggesting that the bird can extend the tongue to access nectar. This study is the first to experimentally confirm bird pollination in the Old World Bignoniaceae. It also indicates aspects that are possibly characteristic of spiderhunter pollination systems, i.e. high degrees of specialization, unusual nectar sugar composition, and floral tubes much longer than bird bills.

Nectar availability and Interactions with Bird Pollinators in Grevillea banksii R. Br. (Proteaceae) in a Restinga Ecosystem in Southeastern Brazil

The mutualistic relationship between plants and their pollinators may influence species’ spatial distribution, richness, and abundance. We assessed the nectar availability and interaction with pollinators in Grevillea banksii (Proteaceae), an alien plant in Brazil. Our objectives were to study the floral biology and the pollination system and identify the pollinators in a G. banksii population in a Restinga ecosystem (sand-coastal-plain) in southeastern Brazil. We described the behavioral parameters of the pollinator-plant interaction and determined how the pollinators contribute to reproduction. Floral biology was analyzed by taking morphometric measurements of the flowers and determining the distribution of their structures. The reproductive system was assessed via different pollination treatments (open pollination, self-pollination, manual selfpollination, manual cross-pollination). The pollinators were identified and their behaviors were assessed by focal observation. Flower morphometrics showed compatibility between the mean calyx length and the beak length of the flower-visiting hummingbirds. All the pollination types tested were efficient, and we identified eight bird species (Chionomesa fimbriata, Eupetomena macroura, Chlorestes notata, Polytmus guainumbi, Florisuga fusca, Coereba flaveola, Cyanerpes cyaneus and Dacnis cayana) that visit G. banksii in the study area. The results show a positive interaction between pollinators and plants, with offers of resources, pollination, and pollen dispersal.

Drivers of sunbird‐plant interactions on Mount Cameroon: Between neutrality and niche‐based processes

AbstractThe drivers behind plant‐pollinator interactions still need to be fully understood. Previous research has suggested that observed interactions result from either neutral interaction between species based on their abundance or from niche‐based processes, which are reflected in the adaptations of both plants and pollinators. Furthermore, the importance of both scenarios can differ depending on seasonal dynamics translated into differences in resource availability. Extensive research has been conducted on New‐World hummingbirds (Trochilidae), whereas much less is known about Old World nectar‐feeding sunbirds (Nectariniidae). Our study aimed to explore whether sunbird‐plant interactions are explained by neutral and/or niche‐based processes. Thus, we tested the effects of abundance, morphology, and nectar sugar content on the observed interactions and the link between sunbirds and plant traits. Moreover, we explored the effects of robbing on these mechanisms. Finally, we investigated the partitioning of bird pollination niches, based on floral traits. We used a dataset of sunbird‐plant interactions collected at Mount Cameroon during two seasons (dry and wet seasons). Our study shows that sunbird‐plant interactions are influenced by both neutral and niche‐based processes. Neutral processes and nectar reward were the main mechanisms underlaying the observed interactions in the dry season. However, as more ornithophilous plants bloom during the rainy season, morphological trait‐matching becomes more important. We found a correlation between bill length and floral tube dimensions and observed niche overlap among the sunbird species. Considering this and other research, we suggest that plant‐pollinator interactions are influenced by a combination of both neutral and niche‐based processes.

Evidence for sunbird pollination in African blood lilies: Haemanthus (Amaryllidaceae)

Flowers arranged in ‘paintbrush’ style inflorescences have evolved in several plant families. Their nectar is often accessible to both birds and insects, meaning that observations and experiments are required in order to establish which animal group pollinates these flowers most effectively. The southern African genus Haemanthus (Amaryllidaceae) includes more than 20 species, all with paintbrush style inflorescences and flower colour that ranges from white to red. Here we examine the contributions of birds versus insects to the pollination of four Haemanthus species varying in flower colour. Direct observations and camera trapping showed that flowers of these Haemanthus species are frequently visited by sunbirds as well as by insects. Bird exclusion experiments revealed that H. coccineus (red flowers) and H. deformis (white flowers) are strongly dependent on sunbirds for pollination, whereas H. humilis subsp. hirsutus (pink flowers) is co-pollinated by insects. All Haemanthus species tested were reliant on pollinators for seed production. Bird pollination appears to be frequent in Haemanthus and is correlated more strongly with large volumes of nectar in inflorescences than it is with a particular flower colour or floral and inflorescence morphology.

Evolution of a non-flying mammal-dependent pollination system in Asian Mucuna (Fabaceae).

Pollinator shifts are often related to speciation in angiosperms, and the relationship between them has been discussed in several plant taxa. Although information on plants pollinated by non-flying mammals in Central and South America and Africa is limited, related research has not been conducted in Asia. Herein, I summarize the available knowledge of pollination in Asian Mucuna (Fabaceae), a genus mainly distributed in the tropics, and discuss the evolution of plants pollinated by non-flying mammals in Asia. Nineteen pollinator species have been recorded and pollination systems have been categorized into four types. An examination of the relationship between Mucuna species and their pollinators from the lineage perspective revealed that all species in M. subgenus Macrocarpa, which are distributed in Asia, are pollinated exclusively by non-flying mammals. Additionally, plants pollinated by non-flying mammals were found to have diverged from bat-pollinated and non-flying mammal-pollinated plants, while plants pollinated by non-flying mammals have evolved multiple times. This is a unique example of evolutionary transition. I hypothesize that the diversification of squirrel species in tropical Asia may have led to the speciation and diversification of Mucuna in Asia. Furthermore, the behavioral and ecological characteristics of bats and birds in Asia differ from the characteristics of those in other regions, implying that Asian Mucuna species do not rely on bat or bird pollinators. The adaptation of floral characteristics to pollinators is not well understood in Asia. Mammal-pollinated plants in Asia may have evolved differently from those in other regions and have unique pollination systems.

Bird and insect pollinators differ in specialization and potential pollination services along disturbance and resource gradients

AbstractCombined studies of the communities and interaction networks of bird and insect pollinators are rare, especially along environmental gradients. Here, we determined how disturbance by fire and variation in sugar resources shape pollinator communities and interactions between plants and their pollinating insects and birds. We recorded insect and bird visits to 21 Protea species across 21 study sites and for 2 years in Fynbos ecosystems in the Western Cape, South Africa. We recorded morphological traits of all pollinator species (41 insect and nine bird species). For each site, we obtained estimates of the time since the last fire (range: 2–25 calendar years) and the Protea nectar sugar amount per hectare (range: 74–62 000 g/ha). We tested how post‐fire age and sugar amount influence the total interaction frequency, species richness and functional diversity of pollinator communities, as well as pollinator specialization (the effective number of plant partners) and potential pollination services (pollination service index) of insects and birds. We found little variation in the total interaction frequency, species richness and functional diversity of insect and bird pollinator communities, but insect species richness increased with post‐fire age. Pollinator specialization and potential pollination services of insects and birds varied differently along the environmental gradients. Bird pollinators visited fewer Protea species at sites with high sugar amount, while there was no such trend for insects. Potential pollination services of insect pollinators to Protea species decreased with increasing post‐fire age and resource amounts, whereas potential pollination services of birds remained constant along the environmental gradients. Despite little changes in pollinator communities, our analyses reveal that insect and bird pollinators differ in their specialization on Protea species and show distinct responses to disturbance and resource gradients. Our comparative study of bird and insect pollinators demonstrates that birds may be able to provide more stable pollination services than insects.

Birds Visiting Flowers of Erythrina suberosa: Their Abundance, Frequency of Visits and Role as Pollinators in a Sub-Tropical Montane Forest of Garhwal Himalaya

Bird pollination is specific among flowering plants which support mostly cross pollination and has been regarded as an important pollination syndrome. Frequency of bird pollinators visiting flowers of the corky coral tree Erythrina suberosa Roxb., was studied in a subtropical montane forest located in Garhwal Himalaya, India. Forty trees were selected randomly in the population and the observations on birds visitors were recorded for 10 days during peak flowering. In total 18 bird species were found visiting flowers of E. suberosa. The bird species belonging to Passeriformes, Piciformes, Psittaciformes, and Cuculiformes were observed most frequently. The highest bird frequency and abundance among flower-visiting birds were recorded for the red-billed blue magpie Urocissa erythrorhyncha (26 ± 3.2 individuals per hour per branch and 60 birds per tree, respectively), whereas the lowest – for the verditer flycatcher Eumyias thalassinus (2 ± 0.02 individuals per hour per branch and 6 individuals per tree, respectively). Majority of the bird species followed bimodal pattern of foraging on nectar in a day (mostly morning and evenings), which is consistent with other studies carried out for other ornithophilous tree species in the Himalayan region. The birds observed in this study are presumed to be pollinators, as the majority of birds foraged on nectar of properly opened flowers oriented upwards; however, possibility of nectar robbing cannot be excluded and requires further investigation in future studies.

Pollen limitation failing reproductive success in selected animal pollinated trees of tropical moist deciduous forest of north-eastern hill region, India

Abstract The aim of this study was to estimate reproductive effort and success in tropical trees and to examine the effect of pollen limitation on reproductive success. Pollen limitation was assessed through pollen supplementation experiment to contrast the open pollination treatment. The taxa selected were Bombax ceiba, Erythrina stricta (ornithophilous trees), Lagerstroemia speciosa, Mesua ferrea and Schima wallichii (entomophilous trees). Index of pollen limitation was highest in Bombax ceiba and Erythrina stricta (both self-incompatible species). The remaining three species were partially self-incompatible with favouring selfing in Lagerstroemia speciosa and Mesua ferrea and supporting outcrossing in Schima wallichii. Therefore, the high index of pollen limitation in Bombax ceiba and Erythrina stricta might be due to the effect of either lacking quality pollen or lacking bird pollinators. All five species produce a large amount of pollen at individual tree level in the order of 108 (Erythrina stricta) to 1010 (Mesua ferrea). Fruit and seed set following pollen supplementation were higher than the open pollination (as control) in all studied species. Pollen limitation in this study is likely associated with the effectiveness of pollinator and their frequency, as all the studied species had produced ample pollen at tree crown level which ultimately leads to pollinator resource limitation in tropical trees.

Using environmental DNA to investigate avian interactions with flowering plants

AbstractAnimal pollination is an important and highly valued ecosystem function and the role of birds as pollinators is increasingly acknowledged. However, such interactions can be challenging to document and often require extensive field programs. Over the last decade, environmental DNA (eDNA) has been analyzed from several different contemporary sample types, such as water, soil, flowers, and air. The applications of these studies include biodiversity monitoring, detection of endangered species, community compositions, and more recently, flower–arthropod interactions. However, it remains unknown whether flower eDNA is applicable to other taxonomic groups interacting with plants, as well as the deposition and degradation of eDNA on flowers. Here, we test whether eDNA from flowers can be used for detecting bird pollinators. In a controlled environment (an aviary with great tits [Parus major]), we show that birds leave significant traces of DNA on the flowers without observed visits (airborne eDNA). We further show that when birds had been in contact with the flowers, DNA concentrations increased to levels significantly higher than airborne background DNA. Subsequently, we sampled five clusters of wild flowers in Papua New Guinea and detected four species of birds, two of which are nectar‐feeders, and one that is an insectivorous species known to visit flowers. These four bird species were regularly seen in the area and caught in mist‐nets in the days prior to sampling of the flowers. In total, 29 bird species were recorded (18 mist‐netted) in the area and of these, eight are nectarivorous. Our quantitative approach suggests that it is possible to distinguish airborne background bird DNA deposited on flowers from actual flower visits of birds in the wild, although this might be highly context‐specific. Our findings are of broad interest within research on ecosystem functioning, biotic interactions, and plant–animal mutualism.

Issue Information

A Orobanche boninsimae and its visitors, Japanese white‐eyes (Zosterops japonicus). Photo by Akihiro Nishimura at Chichijima Island, the Bonin Islands, Japan. Nishimura & Takayama (2023) “First record of potential bird pollination in the holoparasitic genus Orobanche L.” Plant Species Biology, 6–17 https://doi.org/10.1111/1442-1984.12389image

Did early shifts to bird pollination impose constraints on Salvia flower evolution?

A recent article by Kriebel et al. (2020) examines the relationship between floral shape evolution and pollination shifts in Salvia, a plant genus almost worldwide distributed, but particularly diverse in the New World. Kriebel et al. (2020) argue that a major shift to bird pollination at the origin of the subg. Calosphace (~20 million years ago [Mya]) imposed a legacy of constraints, resulting in significant differences in flower morphology between New World and Old World Salvia. However, reanalyses of the data using hidden states to account for the heterogeneity in evolutionary rates do not support an early origin of bird pollination in this group. Instead, bird pollination may have appeared after the arrival of modern hummingbirds to North America (15.5-12 Mya), as in other North American plant clades. The use of more complex models of ancestral state reconstruction into comparative analyses provides a different perspective to explain morphological differences within Salvia. Our results indicate that bird pollination did not impose constraints on corolla shape evolution. Evolutionary constraints in anther connective and style shapes may have arisen at the origin of Calosphace but they were not associated with shifts to hummingbird pollination, being more likely the product of contingent evolution.

High outcrossing rates in a self-compatible and highly aggregated host-generalist mistletoe.

Plants have evolved various strategies to avoid inbreeding, but the mass flowering displayed by many plants predisposes them to within-plant pollen movements and self-pollination. Mistletoes often aggregate at multiple spatial scales. Their bird pollinators often visit several flowers of the same individual and of others on the same host tree. We hypothesized that hermaphroditic mistletoes have self-incompatibility mechanisms that reduce or prevent selfing. Whether their spatial distribution, affected by host specificity, host distribution, and the behaviour of seed dispersers, influences their mating system and population genetic structure remains unclear. We studied how mating system and spatial distribution affect genetic structure in four populations of the host-generalist mistletoe Dendrophthoe pentandra in southwestern China using microsatellite markers and progeny arrays. We also characterized the fine-scale spatial genetic structure among 166 mistletoes from four host trees in one population. Prevalence and intensity of infection both appeared to vary among host species, strongly affecting the degree of aggregation. Host tree size had a strong effect on infection intensity. Surprisingly, manual pollination experiments indicated that D. pentandra is self-compatible, but genetic analyses revealed that outcrossing rates were higher than expected in all four populations (MLTR tm 0.83-1.20, Bayesian tm 0.772-0.952). Spatial genetic structure was associated with distance between host trees but not at shorter scales (within hosts). Our results demonstrate that the combination of bird pollination, bird-mediated seed dispersal, and post-dispersal processes result in outcrossing and maintain relatively high diversity in the presence of biparental inbreeding, despite very high local densities and possible self-compatibility.

First record of potential bird pollination in the holoparasitic genus Orobanche L.

AbstractPollinators play an important role in the reproduction of zoophilous plants. A shift in pollinators has often been observed for oceanic island plants, probably because of the differences in fauna. In this study, we obtained data on pollinator shifts from insects to birds in Orobanche boninsimae (Orobanchaceae), a holoparasitic plant species endemic to the Bonin (Ogasawara) Islands, oceanic islands in the Pacific. We observed pollination and measured seed viability in O. boninsimae and its continental sister species O. coerulescens. We found that two passerine birds, the Japanese white‐eye (Zosterops japonicus) and bulbul (Hypsipetes amaurotis squameiceps), visited the flowers and sucked the nectar of O. boninsimae, while only insects visited those of O. coerulescens. Viable seeds were produced under pollinator‐excluded treatments in the two Orobanche species, indicating that the seeds were produced by automatic self‐pollination and/or apomixis. These results suggest that O. boninsimae may be pollinated by birds and can produce seeds by automatic self‐pollination/apomixis. This is the first record of visitation of the genus Orobanche by birds. Studies of pollination systems in native plants on the Bonin Islands are few compared to those on other oceanic islands, and O. boninsimae may provide a valuable example of pollinator shifts in the Bonin Islands.

Bird pollination

Johnson gives an overview of bird pollinators and the plant species they pollinate.

Latitudinal gradients in biotic interactions: Are cacti pollination systems more specialized in the tropics?

Biotic interactions are said to be more specialized in the tropics, and this was also proposed for the pollination systems of columnar cacti from North America. However, this has not yet been tested for a wider set of cactus species. Here, we use the available information about pollination in the Cactaceae to explore the geographic patterns of this mutualistic interaction, and test if there is a latitudinal gradient in its degree of specialization. We performed a bibliographic search of all publications on the pollination of cacti species and summarized the information to build a database. We used generalized linear models to evaluate if the degree of specialization in cacti pollination systems is affected by latitude, using two different measures: the number of pollinator guilds (functional specialization) and the number of pollinator species (ecological specialization). Our database contained information about the pollination of 148 species. The most frequent pollinator guilds were bees, birds, moths and bats. There was no apparent effect of latitude on the number of guilds that pollinate a cactus species. However, latitude had a small but significant effect on the number of pollinator species that service a given cactus species. Bees are found as pollinators of most cactus species, along a wide latitudinal gradient. Bat and bird pollination is more common in the tropics than in the extra-tropics. The available information suggests that cacti pollination systems are slightly more ecologically specialized in the tropics, but it does not support any trend with regard to functional specialization.

The genetic mechanisms underlying the convergent evolution of pollination syndromes in the Neotropical radiation of Costus L.

Selection together with variation in floral traits can act to mold floral form, often driven by a plant’s predominant or most effective pollinators. To investigate the evolution of traits associated with pollination, we developed a phylogenetic framework for evaluating tempo and mode of pollination shifts across the genus Costus L., known for its evolutionary toggle between traits related to bee and bird pollination. Using a target enrichment approach, we obtained 957 loci for 171 accessions to expand the phylogenetic sampling of Neotropical Costus. In addition, we performed whole genome resequencing for a subset of 20 closely related species with contrasting pollination syndromes. For each of these 20 genomes, a high-quality assembled transcriptome was used as reference for consensus calling of candidate loci hypothesized to be associated with pollination-related traits of interest. To test for the role these candidate genes may play in evolutionary shifts in pollinators, signatures of selection were estimated as dN/dS across the identified candidate loci. We obtained a well-resolved phylogeny for Neotropical Costus despite conflict among gene trees that provide evidence of incomplete lineage sorting and/or reticulation. The overall topology and the network of genome-wide single nucleotide polymorphisms (SNPs) indicate that multiple shifts in pollination strategy have occurred across Costus, while also suggesting the presence of previously undetected signatures of hybridization between distantly related taxa. Traits related to pollination syndromes are strongly correlated and have been gained and lost in concert several times throughout the evolution of the genus. The presence of bract appendages is correlated with two traits associated with defenses against herbivory. Although labellum shape is strongly correlated with overall pollination syndrome, we found no significant impact of labellum shape on diversification rates. Evidence suggests an interplay of pollination success with other selective pressures shaping the evolution of the Costus inflorescence. Although most of the loci used for phylogenetic inference appear to be under purifying selection, many candidate genes associated with functional traits show evidence of being under positive selection. Together these results indicate an interplay of phylogenetic history with adaptive evolution leading to the diversification of pollination-associated traits in Neotropical Costus.