Sugar preferences and digestive efficiency in an opportunistic avian nectarivore, the Dark-capped Bulbul Pycnonotus tricolor

It has recently been recognized that flowers pollinated by generalist opportunistic nectarivores tend to have different nectar properties to those pollinated by specialist nectarivores (including both hummingbirds and specialist passerines). While renewed interest in specialist avian nectarivore sugar preferences and digestive physiology has helped explain the concentrated sucrose-dominated nectar of plants they feed on, there has been little progress in understanding why generalist or occasional nectar-feeding birds tend to be associated with flowers that have dilute hexose-dominated nectar. We examined sugar preferences and assimilation efficiencies over a range of concentrations, and concentration preferences, in Dark-capped Bulbuls Pycnonotus tricolor, one of the more common occasional avian nectarivores in southern Africa. Dark-capped Bulbuls showed significant preference for hexose sugar solutions, irrespective of concentration, when given a choice between hexose and sucrose solutions in equicaloric pair-wise choice tests conducted at five different concentrations (5–25%). This contrasts with results from specialist nectarivore groups which generally show a significant concentration-dependant switch in preference from hexose at low concentrations to sucrose at high concentrations for equicaloric solutions. In addition, Dark-capped Bulbuls showed an unusual lack of preference for solutions of higher sugar concentration when simultaneously offered four solutions varying in concentration from 10 to 25%. Dark-capped bulbuls also showed a unique effect of concentration on sugar assimilation efficiency, assimilating relatively more energy on 5% diets than on 25% diets. Although able to assimilate sucrose effectively, assimilation rates of hexose sugars were marginally higher. These results shed new light on pollination systems involving occasional nectarivores and, in particular, help to explain the prevalence of low concentration hexose-dominated nectars in flowers pollinated by these birds.     

Foraging Blue Tits Parus caeruleus may trade-off calorific value of food items and distance from cover

Capsule Flowers of an invasive plant species are more visited by native birds than flowers of ornithophilous endemic plants.

Aims To describe the bird guild and its behaviour visiting the century plant Agave americana in an insular environment and to determine which factors are affecting visitation rates.

Methods We noted number and species of birds visiting inflorescences on Tenerife, Canary Islands. We used multimodel inference of generalized linear models to analyse the factors affecting the number of visits and the visitor species richness.

Results Eighty-one per cent of inflorescences were visited by eight native bird species. All species fed on nectar and only the Atlantic Canary fed also on pollen. Foraging behaviour varied among species. Visitation rate increased with density and diversity of birds and flower characteristics and decreased through the day. The number of species visiting the inflorescences increased with diversity and density of birds in the surroundings and decreased through the day.

Conclusion The native bird community uses the invasive century plant as a feeding resource at a higher rate than it uses endemic ornithophilous plants. This could have negative effects for the pollination of endemic plants, but positive effects for birds.     

The Macaronesian bird-flower element and its relation to bird and bee opportunists

OLESEN, J. M., 1985. The Macaronesian bird-flower element and its relation to bird and bee opportunists. Canarina canadensis, three Isoplexis species and Lotus berthelotii, among other species, have been supposed to be bird pollinated. None of their flowers reflect UV light and the volume of nectar produced is relatively high with low sugar concentration, indicating bird pollination. This study gives the first report of birds visiting and thus pollinating these species. Sylvia melanocephala, S. conspiciltata and Phylloscopus collybita visited I. canariensis and C. canariensis flowers. Isoplexis sceptrum and Digitalis obscura are regarded as transitional species anthecologically between I. canariensisj I. isabelliana and European Digitalis species.     

Bird–flower interactions in the Macaronesian islands

Aims Several bird‐pollinated or ornithophilous flowers are present on the Macaronesian archipelagos, the Canary Islands and Madeira, but absent from nearby NW Africa and Europe. In Macaronesia, no specialist nectar‐feeding birds are found, but several generalist passerine bird species visit flowers for nectar. Two hypotheses attempt to explain the origin and evolution of ornithophily in the Macaronesian flora. According to ‘the island de novo hypothesis’, bird‐flowers evolved from mainland insect‐pollinated ancestors after island colonization. Alternatively, ancestors of the ornithophilous Macaronesian plant species evolved bird‐flowers before reaching the islands (‘the relict hypothesis’). In this study we first compile information of Macaronesian bird–flower interactions from the literature and our own field observations. Secondly, we discuss the two hypotheses of origin of ornithophily in the light of evidence from recent molecular plant phylogenies, palaeontology, historical biogeography of the African avifauna and flora, and present‐day ecological patterns. Location Madeira and Canary Islands. Results At least eleven endemic Macaronesian plant species from six genera have typical ornithophilous floral traits. These genera are: Canarina and Musschia (Campanulaceae), Isoplexis (Scrophulariaceae), Echium (Boraginaceae), Lotus (Fabaceae) and Lavatera (Malvaceae). These lineages have clear affinities to the Mediterranean region, except for Canarina whose closest relatives grow in East African mountains. Six generalist passerine bird species of Sylvia, Phylloscopus (Sylviidae), Serinus (Fringillidae) and Parus (Paridae) visit this flora for nectar. Main conclusion We suggest that the origin and evolution of ornithophilous traits in these plant species took place mostly in mainland areas prior to island colonization. In Canarina and Lavatera, it is well supported that ornithophily is a relict condition, which originated in mainland areas possibly in association with specialist nectar‐feeding birds. For the remaining plant species except Echium wildpretii bird floral traits probably also are a relict condition. These species may be derived from ancestors, which were visited by specialist nectar‐feeding birds during geological periods when the Mediterranean and the Ethiopian vegetation were intermingled in mainland Africa. Probably, these mainland ancestors went extinct due to severe climatic fluctuations, while their Macaronesian descendants survived in ‘refuge’ on the islands. Finally, the island de novo hypothesis may explain the evolution of a mixed bird/insect‐pollination system in the neo‐endemic red‐flowered Echium wildpretii.     

Bird pollination syndrome is the plant's adaptation to ornithophily,but nectarivorous birds are not so selective

Many tropical plants are pollinated by birds and several bird phylogenetical lineages have specialised to a nectar diet. The long-assumed, intimate ecological and evolutionary relationship between ornithophilous plants and phenotypically specialised nectarivorous birds has nevertheless been questioned in recent decades, where such plant–pollinator interactions have been shown to be highly generalised. In our study, we analysed two extensive interaction datasets: bird–flower and insect–flower interactions, both collected on Mt Cameroon, west-central Africa. We tested if: 1) insects and birds interact with distinct groups of plants; 2) plants with a typical set of ornithophilous floral traits (i.e. bird pollination syndrome) interact mainly with birds; 3) birds favour plants with bird pollination syndrome and; 4) if and how the individual floral traits and plant level nectar production predict bird visitation. Bird-visited plants were typically also visited by insects, while approximately half of the plants were visited by insects only. We confirmed the validity of the bird pollination syndrome hypothesis, as plants with bird-pollination syndrome traits were visited by birds at a higher rate and mostly hosted a lower frequency of visiting insects. However, these ornithophilous plants were not more attractive than the other plants for nectar-feeding birds. Nectar production per plant individual was a better predictor of bird visitation than any other floral trait traditionally related to the bird pollination syndrome. Our study thus demonstrated the highly asymmetrical relationship between ornithophilous plants and nectarivorous birds.     

African sunbirds hover to pollinate an invasive hummingbird-pollinated plant

Why do hummingbirds hover while Old World nectar‐feeding birds perch? A unique opportunity to explore this question is presented by the invasion into Africa of a plant adapted for pollination by hovering hummingbirds. Like other hover‐pollinated plants of the New World, the flowers of the tree tobacco Nicotiana glauca lack perches and are oriented towards open space. We find that Old World nectarivores, especially the malachite sunbird, Nectarinia famosa, hover 80% of the time when taking nectar from these flowers. They hover for up to 30 s, and are able to sustain this hovering lifestyle in an area where native nectar plants are absent. Nicotiana glauca greatly increases the local abundance of sunbirds compared with uninvaded areas. In turn, flowers visited by sunbirds formed significantly more capsules and set significantly more seed than sunbird‐excluded flowers, possibly facilitating the invasion. The results suggest a prominent role for plant, rather than bird traits in determining the occurrence of hover‐pollination, begging the question of why plants adapted for hover pollination do not occur outside the New World.     

The birds and the seeds: opportunistic avian nectarivores enhance reproduction in an endemic montane aloe

Opportunistic bird pollination has become more evident in studies that confirm distinct differences in floral adaptations that attract opportunistic, rather than specialist, bird pollinators. Pollination syndrome studies investigating the effectiveness of different pollinator guilds on reproduction seldom do so by measuring seed viability. We studied pollination in Aloe peglerae, a high altitude endemic succulent of the Magaliesberg mountain range, previously thought to be largely sunbird (specialist) pollinated. Using field observation and pollinator exclusion treatments, i.e. (i) open to all visitors, (ii) bird excluded, and (iii) all visitors excluded, we established that birds contributed significantly more to fruit (2.3–5.6 times) and seed (1.3–1.4 times) set than insect or self-pollination, respectively. Overall, pollination by opportunistic avian nectarivores amplified seed production per aloe ~7 and 10 times compared to insects and self-pollination, respectively. One of three opportunistic nectarivores, the Cape Rock-Thrush (Monticola rupestris), played a significant role in pollination, contributing ~60 % of all probes at inflorescences. The difference in reproductive output between insect visited and visitor excluded flowers was not significant and suggests possible self-pollination in A. peglerae which is particularly unusual in Aloe species. Breeding system experiments would help clarify this. In assessing the effectiveness of pollinator guild on seed viability, we found no differences in percentage seed viability, seed germination or seedling emergence between exclusion treatments. Seed viability and germination were low and variable; however, ~19 % seedling emergence was observed across the treatments. Practically, the net effect of bird pollination may result in 8–12 times more potential seedlings compared to insect and self-pollination respectively. These findings highlight the importance of pollination by opportunistic avian nectarivores in Aloe.     

Honeyeaters and the New Zealand forest flora: The utilisation and profitability of small flowers

New Zealand flowers are frequently considered unspecialised allowing easy access to pollen and nectar by a wide range of visitors. Most conform with a syndrome of insect pollination (entomophily). Pollination of forest flowers by birds has been described for a range of species whose flowers are morphologically ornithophilous. On Kapiti Island and Little Barrier Island, all three species of New Zealand honeyeaters have been described feeding on flowers currently assumed to be entomophilous or where the pollination system is unknown. The persistence and regularity of visits suggests that the birds are obtaining suitable rewards in the form of nectar and could be serving as pollinators. We measured the nectar energetic value from flowers of three ornithophilous and five entomophilous species. Nectar production over 24 hours was highest in ornithophilous species, but the standing crop of nectar overlapped—ornithophilous species: Metrosideros fulgens (standing crop 6.6 J), Metrosideros excelsa (22 J), and Fuchsia excorticata (1.8 J); and entomophilous: Pittosporum crassifolium (23 J), Pseudopanax arboreus (1.5 J), Dysoxylum spectabile (3.7 early flowers -6.7 J late flowers), Pittosporum eugenioides (2.7 J) and Geniostoma rupestre (1.8 J). The entomophilous species present the flowers in aggregation and as result birds can visit a large number flowers per minute. We found that the average estimated nectar consumption rate for all the entomophilous species except G. rupestre was enough to sustain the two smaller New Zealand honeyeaters (hihi energy requirements= 0.12 kJ min(-1), median energy obtained: 0.16 kJ min(-1) D. spectabile—0.57 kJ min(-1) P. crassifolium); bellbird energy requirements = 0.10 kJ min(-1), median energy obtained: 0.14 kJ min(-1) D. spectabile—0.68 kJ min(-1) P. crassifolium). However, we estimate that if the birds are able to selectively forage on the flowers with most nectar, the energetic returns of all species may be sufficient for hihi and bellbird (hihi: 0.18 kJ min(-1) G. rupestre—0.93 kJ min(-1); P. crassifolium; bellbird: 0.12 kJ min(-1) G. rupestre 1.11 kJ min(-1) P. crassifolium). If tui (energy requirements: 0.25 kJ min(-1), forages randomly, only P. crassifolium (0.80 kJ min(- 1)) and D. spectabile late in the season (0.30 kJ min(-1)) provide sufficient returns, but if selective, P. arboreus (0.45 kJ min(-1)) may also suffice. We suggest that because (a) the nectar produced by entomophilous flowers provides sufficient energy to sustain the energetic requirements of birds, and (b) these plants flower in the cooler months when insect activity is reduced, birds might have played a wider role in pollination than previously considered. This finding is of particular importance because the abundance of New Zealand honeyeaters on the mainland has decreased considerably since human colonisation and this could be affecting forest regeneration.     

Ornithophilous canopy species in the Atlantic rain forest of southeastern Brazil

ABSTRACT.   In tropical ecosystems, birds play a relevant role in plant reproduction. Although hummingbirds are regarded as the most important vertebrate pollinators in the Neotropics, the possible role of perching birds as pollinators has been neglected. From 2003 to 2005, we observed 68 species of plants visited by birds in an Atlantic rainforest in southeastern Brazil, including three canopy species: Spirotheca rivieri (Malvaceae, Bombacoidea), Schwartzia brasiliensis (Marcgraviaceae), and Psittacanthus dichrous (Loranthaceae). Flowers of these three species were visited by 15 different species of perching birds and by hummingbirds. The flowers of these three plants are colorful, ranging from deep red or purple to orange. Spirotheca rivieri blooms during the austral winter and Schwartzia brasiliensis during the summer. The flowers of these two species produce copious amounts of dilute nectar in easily accessible structures and both species appear to depend primarily on perching birds as pollinators, with hummingbirds being secondary or minor pollen vectors. The tubular, narrow flowers of P. dichrous are produced during the austral summer and are visited primarily by hummingbirds. Perching birds also visit the flowers, but destroy them. Our results suggest that previous estimates of the number of perching birds that feed on nectar may be too low and that flowers pollinated by perching birds may be more common in the canopy of Neotropical forests than previously thought.     

Efficient avian pollination of Strelitzia reginae outside of South Africa

In its native South Africa, endemic birds pollinate the complex flowers of Strelitzia reginae (bird of paradise) through a highly complex method of pollination. The plant is cultivated worldwide in warm-temperated regions but systematic pollination of the ornithophilous species by local birds has not been reported, and, consequently, seed production is rare outside of South Africa. We found that a member of the New World warblers, Geothlypis trichas, efficiently carried out pollination of S. reginae in southern California, thereby supplementing its typical diet of insects with the energy-rich nectar of S. reginae. Only occasionally, seeds were found in plantings not visited by these birds. The pollinator service provided by the warbler increases seed production in an area outside of South Africa. This could lead to adaptive changes in the exotic species, advance species establishment and persistence and possibly promote invasive behavior in a non-native environment.     

The assembly of plants used as nectar sources by hummingbirds in a Cerrado area of Central Brazil

Studies on hummingbird–plant interactions commonly use a pollination approach emphasizing mutualistic relationships. But floral resources are often used opportunistically by these birds. Plant–pollinator assemblies and pollination sustainability will depend both on the well-adapted plants and other potential floral resources. The Cerrado, Neotropical savannas of Central Brazil, has ca. 7.5 % of its flora supposedly adapted to hummingbird pollination. But detailed information about flowers effectively used by hummingbirds at community level is still lacking. Hence, we recorded all plant species visited by hummingbirds, to determine how these nectariferous flowers were distributed in time and space in different plant formations of a Cerrado area, and also the hummingbird species that visit them. The study was conducted between March 2007 and December 2008 in the Panga Ecological Station. Data regarding flowering phenology, floral morphology and visitation were collected monthly. Forty-six nectariferous species from 39 genera and 17 families were recorded, most with annual flowering dynamics and tubular flowers. But only 21 species had a combination of traits fitting classic ornithophilous syndrome. For the remaining species hummingbird visitation was ascertained from observations at the study site or other sites in the region. Eight hummingbird species occurred in the area and were recorded visiting directly 36 plant species. The study area presented a relatively low number of ornithophilous plants, but a great habitat diversity and many non-ornithophilous plants that hummingbirds used as nectar sources. Therefore, in the studied Cerrado, the diversity of environments and nectariferous plants favour the maintenance of resident and migrant hummingbirds.     

Nectar Production of Calliandra longipedicellata (Fabaceae: Mimosoideae), an Endemic Mexican Shrub with Multiple Potential Pollinators

The nectar resource environment across which nectarivores forage may be patchy and variable. To understand the sources and consequences of such a variation, nectar production was investigated in Calliandra longipedicellata . Nectar was measured once a month throughout a 3-mo winter season in two successive years at three sites. We also conducted diurnal and nocturnal field observations to describe visitation rates of floral visitors, and a pollinator exclusion experiment to evaluate diurnal and nocturnal pollination at the three sites. In all populations, nectar secretion was primarily nocturnal, although flowers produced some nectar during the day. Sugar production per flower varied significantly at both the seasonal and population levels, although nectar production rates and a well-defined afternoon to morning production pattern were consistent across months, populations, and years. Average nectar production rates were high compared to other Calliandra species, and to most hummingbird- or hawkmoth-pollinated plants in the region. Flowers were regularly visited by hawkmoths, bats, hummingbirds and various diurnal insects, and all populations had similar rates of visitation. Nocturnal insects had the highest overall visitation rates (three times as high as those by diurnal insects). Fruit production varied among pollination treatments and populations, and significant differences were found in fruit production when flowers exposed to both diurnal and nocturnal visitation were compared to flowers exposed only to diurnal visitation. Our results and the bright-red staminal filaments of C. longipedicellata indicate lack of specialization for particular pollinators.     

Reproductive biology of Dyckia excelsa Leme (Bromeliaceae): a xerophyte species from ironstone outcrops in central-western Brazil

Dyckia is the third most diverse genus from Pitcairnoideae, pollinated mainly by insects and hummingbirds. In this study, we evaluated demography, reproductive biology and pollination of Dyckia excelsa, a saxicolous species typical of ironstone outcrops. It occurs in groupings composed of genets, ramets and seedlings and bloomed several times, principally in September, with fruiting and dispersing of seeds in November. The anthesis is diurnal and flowers remain open for 2 days. Nectar volume decreased during the anthesis, whereas nectar sugar concentration remained constant. Dyckia excelsa is partially self-compatible and xenogamic; the highest fruit set was observed in the control and cross-pollination treatments; high seed germination rates were observed in all treatments. Flowers have the ornithophilous syndrome, but hummingbirds, bees and butterflies visited them. The occurrence of groupings with multiple inflorescences could contribute to geitonogamy. The high number of flowers during a year supplies food resources and ensures the fidelity of pollinators.     

Evolutionary associations between nectar properties and specificity in bird pollination systems

A long-standing paradigm in biology has been that hummingbirds and passerine birds select for different nectar properties in the plants they pollinate. Here we show that this dichotomy is false and a more useful distinction is that between specialized and generalized bird pollination systems. Flowers adapted for sunbirds, which are specialized passerine nectarivores, have nectar similar to that of hummingbird flowers in terms of volume (approx. 10-30 microl), concentration (approx. 15-25% w/w) and sucrose content (approx. 40-60% of total sugar). In contrast, flowers adapted to generalized bird pollinators are characterized by large volumes (approx. 40-100 microl) of extremely dilute (approx. 8-12%) nectar with minimal sucrose (approx. 0-5%). These differences in nectar traits are highly significant even when statistical analyses are based on phylogenetically separate pairwise comparisons between taxa. We present several hypotheses for the association between nectar properties and specificity in bird pollination systems.     

Intra‐floral resource partitioning between endemic and invasive flower visitors: consequences for pollinator effectiveness

1. Sympatric flower visitor species often partition nectar and pollen and thus affect each other's foraging pattern. Consequently, their pollination service may also be influenced by the presence of other flower visiting species. Ants are solely interested in nectar and frequent flower visitors of some plant species but usually provide no pollination service. Obligate flower visitors such as bees depend on both nectar and pollen and are often more effective pollinators. 2. In Hawaii, we studied the complex interactions between flowers of the endemic tree Metrosideros polymorpha (Myrtaceae) and both, endemic and introduced flower‐visiting insects. The former main‐pollinators of M. polymorpha were birds, which, however, became rare. We evaluated the pollinator effectiveness of endemic and invasive bees and whether it is affected by the type of resource collected and the presence of ants on flowers. 3. Ants were dominant nectar‐consumers that mostly depleted the nectar of visited inflorescences. Accordingly, the visitation frequency, duration, and consequently the pollinator effectiveness of nectar‐foraging honeybees (Apis mellifera) strongly decreased on ant‐visited flowers, whereas pollen‐collecting bees remained largely unaffected by ants. Overall, endemic bees (Hylaeus spp.) were ineffective pollinators. 4. The average net effect of ants on pollination of M. polymorpha was neutral, corresponding to a similar fruit set of ant‐visited and ant‐free inflorescences. 5. Our results suggest that invasive social hymenopterans that often have negative impacts on the Hawaiian flora and fauna may occasionally provide neutral (ants) or even beneficial net effects (honeybees), especially in the absence of native birds.     

Consequences of differences in body mass,wing length and leg morphology for nectar-feeding birds

Nectar-feeding birds are prominent in many parts of the world, and vary with respect to body size. Despite the availability of considerable morphometric data, few concerted efforts have been made to assess the influence of attributes such as mass, wing length and leg morphology upon the speed, acceleration, mode and energetic cost of movement by birds between flowers when foraging for nectar. This review attempts to consolidate and interpret available data and highlight areas where further investigations appear warranted. Australian honeyeaters are generally larger, and American hummingbirds smaller, than Hawaiian honeycreepers and sunbirds of Africa or Asia. Sunbirds, honeyeaters and honeycreepers generally perch while extracting nectar from flowers. Hummingbirds usually hover, apparently because suitable perches close to flowers are lacking, and not because hovering increases the speed at which flowers can be visited. Honeyeaters move from one flower to another at speeds that are at least as great as those for hummingbirds. Most passerine nectarivores need to ingest more nectar per day than hummingbirds in order to maintain energy balance, some species devoting more than 60% of the day to foraging. The major consequence of reduced foraging activity by hummingbirds, which spend only 5–30% of the day in this manner, appears to be male emancipation from nest construction and care of offspring. Large nectarivores have a greater capacity to store surplus food and to fast than smaller birds, and so can take advantage of short-lived peaks in nectar abundance. Nectarivores such as honeyeaters should therefore be favoured by the rapid diurnal changes in nectar availability which are characteristic of many Australian and African habitats. Body mass also determines the likely access to rich sources of nectar through size-related interspecific dominance hierarchies. In all families, larger species tend to monopolize the most rewarding nectar supplies, forcing smaller subordinate species to use poorer, more scattered sources. Within particular species, males usually have longer wings and greater masses than females. These variations imply that the two sexes differ with regard to their foraging ecology, although few supporting data are currently available.     

Reproductive biology of Butea monosperma (Fabaceae)

The reproductive biology encompassing phenology, floral biology, pollination and breeding systems, of Butea monosperma, a beautiful tree of the Indian subcontinent, was investigated in a protected dry, deciduous forest located in New Delhi. Phenological studies indicated that although the species shows a regular flowering season, all trees do not flower every year. Flowers are typically papilionaceous; the stigma is wet papillate and the style is hollow. The flowers show characteristics of bird pollination being large and bright orange-red in colour with copious amounts of nectar, and exhibiting diurnal anthesis. Although the flowers are frequented by as many as seven species of birds belonging to six families, only one species, the purple sunbird (Nectarinia asiatica), is the effective pollinator. The flowers are also pollinated by the three-striped squirrel (Funambulus tristiatus). Unlike other flower visitors, these two pollinators forage the nectar from the open side of the keel (legitimate path) during which pollen grains are deposited on their body parts. After the first visit of a sunbird or a squirrel, virgin flowers showed pollen load on the stigma and developed into fruits. B. monosperma shows a weak form of self-incompatibility. Fruit set following manual self-pollination (5.25 %) was comparable with open-pollination (approx. 5 %) but was significantly lower than manual cross-pollination (22.51 %). This indicates that there is a high degree of geitonogamous pollination in this species, which may lead to a weakening of self-incompatibility as a means of reproductive assurance. The results are analysed in the light of prevailing discussions on specialized vs. generalized pollination systems.     

Termites and the turnover of dead wood in an arid tropical environment

Summary The role of sunbirds (Nectariniidae) in the pollination ecology of Strelitzia nicolai (Musaceae) was studied for one year in a coastal dune forest in Zululand, South Africa. It was found that S. nicolai produced large quantities of low quality nectar (1.74 j/l); that nectar production was highest during the day-time; and that the flowers displayed several characteristics attractive to bird-visitors. The flowers were large, conspicuous and provided the birds with a perch, facilitating easy access to the nectar. Flowers were visited by four species of sunbirds: Olive Sunbird Nectarinia olivacea, Grey Sunbird N. veroxii, Black Sunbird N. amethystina, and Collared Sunbird Anthreptes collaris. Sunbirds visited the flowers throughout the year, and apparently cued into changes in the flower angle as an indication of nectar flow rates. Sunbirds perched on the flowers in a manner which effected pollination, the pollen being transferred to the stigma via the birds' feet. Besides the sunbirds, there were other visitors (bushbabies, monkeys and insects) to the flowers, but they did not visit the flowers frequently nor did they appear to be significant pollinators. The high seed set of S. nicolai in the study area attests to the efficacy of the sunbirds as pollinators.     

Ornithophilie auf den Canarischen Inseln

(1) On the Canary Islands and Madeira typical bird-flowers occur in at least twelve species of six genera, although true flower-birds are absent. This inconsistency is in part elucidated by field observations on exotic and wild plants of Tenerife. —(2) In the Botanical Garden of Orotava it could be observed that various ornithophilous plants, which were introduced there, were visited by indigenous birds for nectar and in one case (Orthostemon) for food tissue. Of the three bird species involved, an endemic race of Chiffchaff (Phylloscopus collybita) and resident Blackcaps (Sylvia atricapilla) exploit, and pollinate, flowers legitimously, while the Wild Canary (Serinus canaria) is predominantly a destructive nectar robber. —(3) The insular Chiffchaff also proved to be a regular pollinator in the wild, at least ofCanarina canariensis andIsoplexis canariensis, two ornithophilous paleoendemics. Ornithophily, thus, is naturally practised on the island, though by birds basically insectivorous. —(4) A list of Macaronesian plants bearing the more or less complete ornithophilous syndrome is presented, including newly recognizedTeucrium heterophyllum andScrophularia calliantha. —(5) On biogeographical and faunistic grounds it is presumed that the modern visitors of Canarian bird flowers are secondary rather than the original partners of the continental tertiary flora in which these plants originated. Palearctic immigrants, when becoming resident on the islands during and since the Pleistocene, adopted facultative nectar feeding, entering an orphaned food niche. Casual flower visits in Europe suggest a certain predisposition of the Chiffchaff and the Blackcap for the exploitation of flowers. — (6) The ability of unspecialized birds to acquire nectardrinking spontaneously and to pass this habit on to their offspring, is demonstrated by a population of Tree Sparrows (Passer montanus) which have visited ornithophilousKniphofia (Liliaceae) in Berlin for several years.

     

The Bizarre Inflorescence of Norantea brasiliensis (Marcgraviaceae): Visits of Hovering and Perching Birds1

The pollination biology of Norantea brasiliensis (Marcgraviaceae) was studied in the rain forest of southeastern Brazil. This plant presents bizarre, brush-type racemous inflorescences bearing numerous flowers and extrafloral cup-shaped nectaries. Flower anthesis is diurnal, nectar production is continuous and copious, and the sticky pollen is readily removed by visitors during the first morning hours. The ruby-coloured inflorescences were visited by eight species of hummingbirds (Trochilidae), and 10 species of passerine birds (three Coerebidae and seven Thraupidae). Hummingbirds hovered while probing for nectar and touched flowers occasionally, whereas passerine birds perched and made contact with flowers habitually. Due to differences in flower-visiting and general foraging behaviour, perching birds act as better pollen vectors than hovering birds. The inflorescence of Norantea brasiliensis seems well fitted for pollination by passerine birds, and the hexose-dominated nectar supports this idea. Pollination syndrome trends within Marcgraviaceae may stem from insect-pollinated, condensed and spike-like inflorescences which would give rise to bird-pollinated, brush-type inflorescences. From the same basic condensed inflorescence, bat-pollinated umbelliform inflorescence may be derived from bird-pollinated, pendulous and corymb-like inflorescences. These postulated inflorescence types are found among the extant species of Marcgraviaceae.