Generalist birds outperform specialist sunbirds as pollinators of an African Aloe.

Bird pollination systems are dominated by specialist nectarivores, such as hummingbirds in the Americas and sunbirds in Africa. Opportunistic (generalist) avian nectarivores such as orioles, weavers and bulbuls have also been implicated as plant pollinators, but their effectiveness as agents of pollen transfer is poorly known. Here, we compare the single-visit effectiveness of specialist and opportunistic avian nectarivores as pollinators of Aloe ferox, a plant that relies almost exclusively on birds for seed production. We found that the number of pollen grains on stigmas of flowers receiving single visits by opportunistic avian nectarivores was approximately threefold greater than on those receiving single visits by specialist sunbirds and about twofold greater than on those that received single visits by honeybees. The number of pollen grains on stigmas of flowers visited by sunbirds was similar to that on stigmas of unvisited flowers. These results show that opportunistic birds are highly effective pollinators of A. ferox, supporting the idea that some plants are specialized for pollination by opportunistic birds.

Genomic data resolve gene tree discordance in spiderhunters (Nectariniidae, Arachnothera).

Reduced representation genomic sequencing methods efficiently gather sequence data from thousands of loci throughout the genome. These data can be used to test previous phylogenetic hypotheses produced from limited numbers of mitochondrial and nuclear loci that often reveal intriguing, but conflicting, results. In this paper, we use phylogenomic data to revisit recent molecular phylogenetic work that clarified many taxonomic relationships within spiderhunters, but also questioned the monophyly of this distinctive genus of sunbirds (AVES: Nectariniidae; Arachnothera). DNA sequence data were produced by target-capture sequencing of ultraconserved elements (UCEs) to infer the evolutionary history of 11 species of Arachnothera and six outgroups, including the Purple-naped Sunbird (Hypogramma hypogrammicum), which previous work suggested might lie within Arachnothera. Although we recovered many different gene tree topologies, concatenated and coalescent methods of analysis converged on a species tree that strongly supports the monophyly of Arachnothera, with Hypogramma as its sister taxon.

Do sunbirds use taste to decide how much to drink?

Nectarivorous birds typically consume smaller meals of more concentrated than of less concentrated sugar solutions. It is not clear, however, whether they use taste to decide how much to consume or whether they base this decision on post-ingestive feedback. Taste, a cue to nectar concentration, is available to nectarivores during ingestion whereas post-ingestive information about resource quality becomes available only after a meal. When conditions are variable, we would expect nectarivorous birds to base their decisions on how much to consume on taste, as post-ingestive feedback from previous meals would not be a reliable cue to current resource quality. Here, we tested whether white-bellied sunbirds (Cinnyris talatala), foraging from an array of artificial flowers, use taste to decide how much to consume per meal when nectar concentration is highly variable: they did not. Instead, how much they chose to consume per meal appeared to depend on the energy intake at the previous meal, that is how hungry they were. Our birds did, however, appear to use taste to decide how much to consume per flower visited within a meal. Unexpectedly, some individuals preferred to consume more from flowers with lower concentration rewards and some preferred to do the opposite. We draw attention to the fact that many studies perhaps misleadingly claim that birds use sweet taste to inform their foraging decisions, as they analyse mean data for multiple meals over which post-ingestive feedback will have become available rather than data for individual meals when only sensory information is available. We discuss how conflicting foraging rules could explain why sunbirds do not use sweet taste to inform their meal size decisions.

Drinking problems on a 'simple' diet: physiological convergence in nectar-feeding birds.

Regulation of energy and water are by necessity closely linked in avian nectarivores, because the easily available sugars in nectar are accompanied by an excess of water but few electrolytes. In general, there is convergence in morphology and physiology between three main lineages of avian nectarivores that have evolved on different continents - the hummingbirds, sunbirds and honeyeaters. These birds show similar dependence of sugar preferences on nectar concentration, high intestinal sucrase activity and rapid absorption of hexoses via mediated and paracellular routes. There are differences, however, in how these lineages deal with energy challenges, as well as processing the large volumes of preformed water ingested in nectar. While hummingbirds rely on varying renal water reabsorption, the passerine nectarivores modulate intestinal water absorption during water loading, thus reducing the impact on the kidneys. Hummingbirds do not generally cope with salt loading, and have renal morphology consistent with their ability to produce copious dilute urine; by contrast, as well as being able to deal with dilute diets, honeyeaters and sunbirds are more than capable of dealing with moderately high levels of added electrolytes. And finally, in response to energy challenge, hummingbirds readily resort to torpor, while the passerines show renal and digestive responses that allow them to deal with short-term fasts and rapidly restore energy balance without using torpor. In conclusion, sunbirds and honeyeaters demonstrate a degree of physiological plasticity in dealing with digestive and renal challenges of their nectar diet, while hummingbirds appear to be more constrained by this diet.

Floral colour variation of Nicotiana glauca in native and non-native ranges: Testing the role of pollinators' perception and abiotic factors.

Invasive plants displaying disparate pollination environments and abiotic conditions in native and non-native ranges provide ideal systems to test the role of different ecological factors driving flower colour variation. We quantified corolla reflectance of the ornithophilous South American Nicotiana glauca in native populations, where plants are pollinated by hummingbirds, and in populations from two invaded regions: South Africa, where plants are pollinated by sunbirds, and the Balearic island of Mallorca, where plants reproduce by selfing. Using visual modelling we examined how corolla reflectance could be perceived by floral visitors present in each region. Through Mantel tests we assessed a possible association between flower colour and different abiotic factors. Corolla reflectance variation (mainly along medium to long wavelengths, i.e. human green-yellow to red colours) was greater among studied regions than within them. Flower colour was more similar between South America and South Africa, which share birds as pollinators. Within invaded regions, corolla reflectance variation was lower in South Africa, where populations could not be distinguished from each other by sunbirds, than in Spain, where populations could be distinguished from each other by their occasional visitors. Differences in corolla colour among populations were partially associated with differences in temperature. Our findings suggest that shifts in flower colour of N. glauca across native and invaded ranges could be shaped by changes in both pollination environment and climatic factors. This is the first study on plant invasions considering visual perception of different pollinators and abiotic drivers of flower colour variation.

An invasive alien Proteaceae lures some, but not all nectar-feeding bird pollinators away from native Proteaceae in South African fynbos.

Invasive alien plants often influence pollinator visitation to native plants when sharing pollinator guilds. It is of conservation concern when the invasive alien plant is characterized by floral resources that attract pollinators, thereby reducing the reproductive success of native species. This is well studied for insects, but whether the same is true for bird pollinators is largely unknown. We address this by considering the impact of an invasive alien plant (Banksia speciosa) on visitation rates of nectar-feeding bird pollinators to native Protea compacta in the Cape Floristic Region of South Africa. We determined bird pollinator visitation rate to B. speciosa and P. compacta over 21 h of observation at three sites. We also quantified how visitation rate influenced reproductive success of both study species through different breeding experiments. Sugarbird visitation to P. compacta was significantly lower in the presence of B. speciosa, while there was no effect for sunbirds as they mostly avoided B. speciosa. Protea compacta had higher nectar volumes and sucrose per flower than B. speciosa. Sucrose per hectare was higher in all P. compacta plots compared to B. speciosa. Neither study species is pollen limited and they are self-compatible to some extent. But pollinator visitation enhanced seed production in both species. We show here that the invasive alien B. speciosa flowers attract sugarbirds - but not sunbirds - away from native P. compacta. The long-term effect of reduced pollinator visitation may reduce the fitness of P. compacta, but the long-term demographic impact is unknown and would require further study.

Differences in Nectar Traits between Ornithophilous and Entomophilous Plants on Mount Cameroon.

Despite a growing number of studies, the role of pollinators as a selection agent for nectar traits remains unclear. Moreover, the lack of data from some biogeographic regions prohibits us from determining their general importance and global patterns. We analyzed nectar carbohydrate traits and determined the main pollinators of 66 plant species in the tropical forests of Mount Cameroon (tropical West Africa). The measured nectar traits included total sugar amounts and proportions of sucrose and hexoses (i.e., glucose and fructose). We report the nectar properties for plants visited by five pollinator groups (bees, butterflies, moths, hoverflies, and specialized birds). Our results indicate that, rather than specific evolution in each of the five plant groups, there was a unique nectar-trait evolution in plants pollinated by specialized birds. The ornithophilous plants had a higher proportion of sucrose and produced larger sugar amounts than the plants pollinated by insects. We also demonstrated a significant phylogenetic signal in the nectar properties in some lineages of the studied plants.

Comparison of the ecology and evolution of plants with a generalist bird pollination system between continents and islands worldwide.

Thousands of plant species worldwide are dependent on birds for pollination. While the ecology and evolution of interactions between specialist nectarivorous birds and the plants they pollinate is relatively well understood, very little is known on pollination by generalist birds. The flower characters of this pollination syndrome are clearly defined but the geographical distribution patterns, habitat preferences and ecological factors driving the evolution of generalist-bird-pollinated plant species have never been analysed. Herein I provide an overview, compare the distribution of character states for plants growing on continents with those occurring on oceanic islands and discuss the environmental factors driving the evolution of both groups. The ecological niches of generalist-bird-pollinated plant species differ: on continents these plants mainly occur in habitats with pronounced climatic seasonality whereas on islands generalist-bird-pollinated plant species mainly occur in evergreen forests. Further, on continents generalist-bird-pollinated plant species are mostly shrubs and other large woody species producing numerous flowers with a self-incompatible reproductive system, while on islands they are mostly small shrubs producing fewer flowers and are self-compatible. This difference in character states indicates that diverging ecological factors are likely to have driven the evolution of these groups: on continents, plants that evolved generalist bird pollination escape from pollinator groups that tend to maintain self-pollination by installing feeding territories in single flowering trees or shrubs, such as social bees or specialist nectarivorous birds. This pattern is more pronounced in the New compared to the Old World. By contrast, on islands, plants evolved generalist bird pollination as an adaptation to birds as a reliable pollinator group, a pattern previously known from plants pollinated by specialist nectarivorous birds in tropical mountain ranges. Additionally, I discuss the evolutionary origins of bird pollination systems in comparison to systems involving specialist nectarivorous birds and reconstruct the bird pollination system of Hawaii, which may represent an intermediate between a specialist and generalist bird pollination system. I also discuss the interesting case of Australia, where it is difficult to distinguish between specialist and generalist bird pollination systems.