Native birds and insects,and introduced honey bees visiting Echium wildpretii (Boraginaceae) in the Canary Islands

In this paper, we report observations of flower visitors of the endemic Echium wildpretii in Tenerife, Canary Islands. This plant inhabits the high altitudinal sub-alpine zone, which is characterized by a harsh climate, low species diversity and a short growing season. Echium wildpretii is a monocarpic perennial, producing a 2–3 m column-shaped, red-flowered, nectar-rich inflorescence. Although these floral traits have previously been suggested as being typical of ornithophilous flowers, this is the first study reporting observations of native birds (Phylloscopus collybita and Serinus canarius) in addition to insects visiting the flowers for nectar. The purposes of this study were firstly to investigate levels of visitation by native birds, native insects, and introduced honey bees. Secondly, we studied the influence of floral display (plant height and number of flowers), nearest neighbours (distance and size) and local vegetation structure on visitation rate. Finally, we discuss the evolution of ornithophily in an otherwise entomophilous plant lineage. We found that the level of bird visitation was relatively high early in the flowering season, but decreased in mid/late season, while the opposite pattern was found for introduced honey bees. For native insects, the frequency of visits was similar in early and late season. Bird visits were correlated with floral display. In the early season, visitation rates of honey bees and the two most common native bee species were correlated with size of the plant or its nearest neighbours, consistent with preference patterns for larger resource patches. Since only insects visit the flowers of other species in the Echium clade, E. wildpretii appears to have evolved from a truly insect-pollinated lineage.     

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.

     

Effectiveness of short‐tongued bees as pollinators of apparently ornithophilous New Zealand mistletoes

Abstract The flowers of two species of threatened New Zealand mistletoes (Peraxilla tetrapetala and Peraxilla colensoi, Loranthaceae) have explosive buds that do not open unless force is applied by birds or two species of native short‐tongued bees. Opened flowers are visited by a variety of birds and insects. Although both species of Peraxilla conform to a pollination syndrome of ornithophily, bees may be effective alternative pollinators. We investigated the effectiveness of bees and birds as pollinators of P. colensoi at one site and P. tetrapetala at two sites in the South Island. Bees and other insects outnumbered birds as flower visitors at all three sites. By excluding birds with wire cages, we showed that two bee species regularly open flowers of P. tetrapetala, but only rarely open flowers of P. colensoi. Few pollen grains were deposited when either birds or bees opened buds, so opening buds was not by itself sufficient for adequate pollination. Instead, pollen continued to accumulate over the next 6 or 7 days, even inside cages that excluded birds. Both populations of P. tetrapetala were regularly pollen‐limited, but in different ways. At Ohau, opened flowers gained enough pollen to produce seeds, but many buds were not opened and hence failed to set seed. In contrast, at Craigieburn, nearly all buds were opened, but many of these did not receive enough pollen. These results demonstrate that native bees can partially replace birds as pollinators of mistletoes, despite their apparent ornithophilous syndrome. Ongoing reductions in New Zealand forest bird numbers means that the service bees provide may be important for the long‐term future of these plants.     

Main sugar composition of floral nectar in three species groups of Scrophularia (Scrophulariaceae) with different principal pollinators

In some angiosperm groups, a parallelism between nectar traits and pollination syndromes has been demonstrated, whereas in others there is not such relationship and it has been explained as due to phylogenetic constraints. However, nectar trait information remains scarce for many plant groups. This paper focuses on three groups of Scrophularia species, with different flower sizes and principal pollinators, to find out whether nectar sugar composition is determined by pollinator type or reflects taxonomic affinities. Since the species we examined have protogynous flowers, and gender bias in nectar sugar composition has been noted in few plant groups, we also investigated whether sexual phase influenced Scrophularia nectar composition. The sugar composition was found to be similar in all species, having high‐sucrose nectar, except for the Macaronesian Scrophularia calliantha, which was the only species with balanced nectar; this last kind of nectar could be associated with the high interaction rates observed between S. calliantha and passerine birds. The nectar sugar composition (high in sucrose) was unrelated to the principal pollinator group, and could instead be considered a conservative taxonomic trait. No gender bias was observed between functionally female and male flowers for nectar volume or concentration. However, sexual phase significantly affected sucrose percentage in the largest‐flowered species, where the female phase flowers had higher sucrose percentages than the male phase flowers.     

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.     

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.     

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.     

Covariation of flower traits and bird pollinator assemblages among populations of Kniphofia linearifolia (Asphodelaceae)

Globally, bird-pollinated plants can be separated into two groups, one consisting of species pollinated by specialist nectarivores, and the other of plants pollinated by occasional nectarivores. There are marked differences in nectar properties among the two groups, implying that there has been pollinator-mediated selection on these traits. This raises the possibility that variation in bird assemblages among populations of a plant species could lead to the evolution of intraspecific variation in floral traits. We examined this hypothesis in Kniphofia linearifolia, a common and widespread plant in southern Africa. Although bees are common visitors to flowers of this species, exclusion of birds from inflorescences led to significant reductions in seed set, indicating that the species is primarily bird-pollinated. We showed that bird pollinator assemblages differ markedly between five different populations of K. linearifolia, and that variation in flower morphology and nectar properties between these populations are associated with the dominant guild of bird visitors at each population. We identified two distinct morphotypes, based on corolla length, nectar volume and nectar concentration, which reflect the bird assemblages found in each type. Further work is needed to establish if a natural geographic mosaic of bird assemblages are the ultimate cause of differentiation in floral traits in this species.     

SEASONAL CHANGES IN NECTAR-FEEDING BY BIRDS AT ZARIA, NIGERIA

Nectar-feeding of birds was recorded during a phenological study of the woody vegetation at Zaria in the Northern Guinea Savanna of Nigeria. Throughout the dry season when it is in a non-breeding state, the Scarlet-breasted Sunbird Nectarinia senegalensis concentrates on the ornithophilous plants which flower in an overlapping sequence, and exploits the smaller-flowered species less frequently. As the flowering of the ornithophilous species decreases at the end of the dry season, the range of other plants exploited increases but, in the early rains as the flowering of the indigenous plants tails off, the sunbird switches to ornamental exotics and the indigenous, shrubby parasite, Tapinanthus globiferus. The bird is more conspicuously insectivorous during the wet season when it breeds. The Pygmy Sunbird Anthreptes platura is a breeding, dry-season visitor which exploits a somewhat different range of small-flowered, mainly entomophilous species and some exotics but also takes nectar from those ornithophilous species it can exploit, as well as the chiropterophilous Parkia clappertoniana. The bulk of the breeding population leaves the district before the flowering of the indigenous plant tails off. The five other species of sunbird recorded in the district are either very rare residents or more numerous wet-season visitors and passage migrants for which records of nectar-feeding are too few to draw conclusions about seasonal changes but the wet-season visitors appear to rely on the garden exotics in the comparative absence of flowering indigenous plants. Nectar-feeding on Bombax costatum and Parkia clappertoniana, where nectar is readily accessible and present in relatively large quantities, was noted for IS species of non-nectariniid birds, some of which were regular visitors. Some examples of flower consumption probably related to nectar-feeding are also mentioned.     

An Assemblage of Hummingbird-pollinated Flowers in a Montane Forest in Southeastern Brazil

Relationships between ornithophilous flowers and hummingbirds have been little studied in southern South America, where hummingbird species richness is low. We studied an ornithophilous flower assemblage and the hummingbird pollinators in a montane forest in southeastern Brazil. Twenty-three native hummingbird-pollinated plant species in 21 genera and 14 families were observed. Bromeliaceae, Fabaceae, Gesneriaceae, and Lobeliaceae are represented by more than one species within the assemblage. Flower shapes vary from narrow tube to bowl-shape, but tubular flowers prevail. The variety of flower shapes and sizes results in diverse pollen placement on the body parts of hummingbird visitors, although pollen is deposited mostly on the bill. Sugar concentration in nectar averages 22.1%, and nectar volume per flower averages 16.9 μl. The plant populations bloom for one month to year-round, and their flowering approaches the steady-state pattern. Four flower subsets may be defined within the assemblage, each subset related to the bill size and foraging habits of the most frequent bird visitor. Of the six species of hummingbirds recorded at the study site, four are common and largely resident. The four hummingbirds differ in bill size, body mass, and favoured foraging sites, attributes which reflect their favoured flower subsets. One hermit and one trochiline hummingbird share most of the flower species they use, these two birds being the major pollinators within the flower assemblage. This montane forest community may be viewed as medium-rich in ornithophilous flower species and poor in hummingbird species.     

Mixed hummingbird: Long‐proboscid‐fly pollination in ‘ornithophilous’Embothrium coccineum (Proteaceae) along a rainfall gradient in Patagonia,Argentina

Abstract The pollination ecology of eight populations of the tree Embothrium coccineum was studied along a steep rainfall gradient in NW Patagonia, Argentina. The showy red flowers suggest an ornithophilous pollination syndrome and they have been reported to attract hummingbirds in Argentina and hummingbirds and passerines in Chile. At each population, flower visitors were recorded and floral rewards were analysed. We found a highly significant increase in nectar concentration towards the drier end of the gradient, but this change was not related to the turnover of species in the flower‐visitor assemblage of E. coccineum. In addition to the hummingbird Sephanoides sephaniodes (Green‐Backed Firecrown, Trochilidae) which is widespread throughout the temperate forest at this latitude, other species seem to be locally important as pollinators of E. coccineum in some sites in Argentina (e.g. two long‐tongued tanglewing flies (Nemestrinidae) of the genus Trichophthalma). The long‐dated occurrence of tanglewing flies in South America, relative to the more modern hummingbirds, suggests that ornithophily may be a derived character in E. coccineum, the ancestral condition being pollination by Nemestrinidae.     

Flower visitation by birds in Europe

Most flowering plants depend on animal pollination. Several animal groups, including many birds, have specialized in exploiting floral nectar, while simultaneously pollinating the flowers they visit. These specialized pollinators are present in all continents except Europe and Antarctica, and thus, insects are often considered the only ecologically relevant pollinators in Europe. Nevertheless, generalist birds are also known to visit flowers, and several reports of flower visitation by birds in this continent prompted us to review available information in order to estimate its prevalence. We retrieved reports of flower–bird interactions from 62 publications. Forty‐six bird species visited the flowers of 95 plant species, 26 of these being exotic to Europe, yielding a total of 243 specific interactions. The ecological importance of bird–flower visitation in Europe is still unknown, particularly in terms of plant reproductive output, but effective pollination has been confirmed for several native and exotic plant species. We suggest nectar and pollen to be important food resources for several bird species, especially tits Cyanistes and Sylvia and Phylloscopus warblers during winter and spring. The prevalence of bird flower‐visitation, and thus potential bird pollination, is slightly more common in the Mediterranean basin, which is a stopover to many migrant bird species, which might actually increase their effectiveness as pollinators by promoting long‐distance pollen flow. We argue that research on bird pollination in Europe deserves further attention to explore its ecological and evolutionary relevance.     

AND IF ENGLER WAS NOT COMPLETELY WRONG? EVIDENCE FOR MULTIPLE EVOLUTIONARY ORIGINS IN THE MOSS FLORA OF MACARONESIA

The Macaronesian endemic flora has traditionally been interpreted as a relict of a subtropical element that spanned across Europe in the Tertiary. This hypothesis is revisited in the moss subfamily Helicodontioideae based on molecular divergence estimates derived from two independent calibration techniques either employing fossil evidence or using an Monte Carlo Markov Chain (MCMC) to sample absolute rates of nucleotide substitution from a prior distribution encompassing a wide range of rates documented across land plants. Both analyses suggest that the monotypic Madeiran endemic genus Hedenasiastrum diverged of other Helicodontioideae about 40 million years, that is, well before Macaronesian archipelagos actually emerged, in agreement with the relict hypothesis. Hedenasiastrum is characterized by a plesiomorphic morphology, which is suggestive of a complete morphological stasis over 40 million years. Macaronesian endemic Rhynchostegiella species, whose polyphyletic origin involves multiple colonization events, evolved much more recently, and yet accumulated many more morphological novelties than H. percurrens. The Macaronesian moss flora thus appears as a complex mix of ancient relicts and more recently dispersed, fast‐evolving taxa.     

Floral resources and hummingbirds on an island of flooded forest in Central Brazil

Nectar availability for hummingbirds was studied for two and a half years in an island of flooded forest in Uberlândia, MG, in the Cerrado region, the Neotropical savanna areas in Central Brazil. Flowering individuals of ten plant species from eight families that offered resources potentially used by hummingbirds were recorded along transects crisscrossing the area, which were surveyed every other week. The number of flowers was estimated for each plant. The volume and concentration of nectar were measured for a sample of flowers of each species, and the data were used to estimate the caloric content available per flower and the general resource availability in the area throughout the study period. Focal observations of hummingbird visits were carried out on blooming individuals of all plant species. Hummingbird flowers occurred throughout the study period between January 2001 and June 2003. Only three of the studied plants did not present clear ornithophilous features. Six hummingbird species were recorded in the area, but total nectar availability in the area was seldom sufficient for the daily energy requirements of a single bird. Therefore, it can be inferred from the study that these hummingbirds need to move among the mosaic of different plant physiognomies in the region in order to obtain enough energy for their survival. These movements may also contribute to increasing gene flow and regeneration of plant populations isolated in forest patches as the one studied here.     

Response of avian nectarivores to the flowering of Aloe marlothii : a nectar oasis during dry South African winters

In southern Africa, Aloe marlothii flowers during the dry winter season and offers copious dilute nectar to a variety of birds. Avian abundance and community composition were monitored at an A. marlothii forest at Suikerbosrand Nature Reserve, South Africa. Sampling occurred during two summer months (February–March) when no flowers were present, and six months (May–October) that spanned the winter flowering. We hypothesized that an influx of occasional nectarivores to the A. marlothii forest during flowering would lead to significant changes in the avian community. Overall bird abundance increased 2–3 fold at the peak of nectar availability (August). We recorded 38 bird species, of 83 species detected during transects, feeding on A. marlothii nectar; this diverse assemblage of birds belonged to 19 families, including Lybiidae, Coliidae, Pycnonotidae, Sylviidae, Cisticolidae, Muscicapidae, Sturnidae, Ploceidae and Fringillidae. Surprisingly, only two species of sunbird (Nectariniidae) were observed feeding on A. marlothii nectar, and both occurred in low abundance. We predicted that competition for nectar resources would be high, but few aggressive inter- and intra-specific interactions occurred between birds while feeding on inflorescences. During peak flowering, insect feeders (insectivores, omnivores, nectarivores) fed on nectar during the cold morning when insect activity was low, whilst non-insect feeders (frugivores and granivores) fed on nectar in the middle of the day. Our study highlights the importance of A. marlothii nectar as a seasonal food and water source for a diverse assemblage of occasional nectarivores.     

Relationships among nectar‐dwelling yeasts,flowers and ants: patterns and incidence on nectar traits

Nectar‐dwelling yeasts are emerging as widely distributed organisms playing a potentially significant and barely unexplored ecological role in plant pollinator mutualisms. Previous efforts at understanding nectar–pollinator–yeast interactions have focused on bee‐pollinated plants, while the importance of nectarivorous ants as vectors for yeast dispersal remains unexplored so far. Here we assess the abundance and composition of the nectar fungal microbiota of the ant‐pollinated plant Cytinus hypocistis, study whether yeast transmission is coupled with ant visitation, and discern whether ant‐ transported yeasts promote changes in nectar characteristics. Our results show that a high percentage of flowers (77%) and plants (94%) contained yeasts, with yeast cell density in nectar reaching up to 6.2 × 10⁴ cells mm^?3, being the highest densities associated with the presence of the nectar‐specialist yeast Metschnikowia reukaufii. The establishment of fungal microbiota in nectar required flower visitation by ants, with 70% of yeast species transported by them being also detected in nectar. Ant‐vectored yeasts diminished the nutritional quality of nectar, with flowers exposed to pollinators and yeasts containing significantly lower nectar sugar concentration than virgin flowers (13.4% and 22.8%, respectively). Nectar of flowers that harbored M. reukaufii showed the lowest quality, with nectar concentration declining significantly with increasing yeast density. Additionally, yeasts modified patterns of interpopulation variation in nectar traits, homo genizing differences between populations in some nectar attributes. We show for the first time that the outcome of the tripartite pollinator–flower–yeast interaction is highly dependent on the identity and inherent properties of the participants, even to the extent of influencing the species composition of this ternary system, and can be mediated by ecological characteristics of plant populations. Through their influence on plant functional traits, yeasts have the potential to alter nectar consumption, pollinator foraging behavior and ultimately plant reproduction.     

Species‐specific responses to combined water stress and increasing temperatures in two bee‐pollinated congeners (Echium,Boraginaceae)

1. Water stress and increasing temperatures are two main constraints faced by plants in the context of climate change. These constraints affect plant physiology and morphology, including phenology, floral traits, and nectar rewards, thus altering plant–pollinator interactions.
2. We compared the abiotic stress responses of two bee‐pollinated Boraginaceae species, Echium plantagineum, an annual, and Echium vulgare, a biennial. Plants were grown for 5 weeks during their flowering period under two watering regimes (well‐watered and water‐stressed) and three temperature regimes (21, 24, 27°C).
3. We measured physiological traits linked to photosynthesis (chlorophyll content, stomatal conductance, and water use efficiency), and vegetative (leaf number and growth rate) and floral (e.g., flower number, phenology, floral morphology, and nectar production) traits.
4. The physiological and morphological traits of both species were affected by the water and temperature stresses, although the effects were greater for the annual species. Both stresses negatively affected floral traits, accelerating flower phenology, decreasing flower size, and, for the annual species, decreasing nectar rewards. In both species, the number of flowers was reduced by 22%–45% under water stress, limiting the total amount of floral rewards.
5. Under water stress and increasing temperatures, which mimic the effects of climate change, floral traits and resources of bee‐pollinated species are affected and can lead to disruptions of pollination and reproductive success.

     

The relative importance of birds and insects as pollinators of the New Zealand flora

Native birds may have been underestimated as pollinators of the New Zealand flora due to their early decline in abundance and diversity on the mainland. This paper reconsiders the relative importance of birds and insects as pollinators to eight native flowering plants, representing a range of pollination syndromes, on two offshore island refuges. Experimental manipulations were made on five of these plant species to assess the relative effectiveness of bird and insect visitors as pollinators. In addition, foraging behaviour and the respective morphologies of flowers and visitors were measured at all eight plants to identify the main pollinators. The experimental measures showed that percentage fruit set was significantly higher in flowers exposed to birds than flowers from which birds were excluded in all manipulated plants. The observational measures revealed that for six of the flowering species (Sophora microphylla, Vitex lucens, Pittosporum crassifolium, Pittosporum umbellatum, Pseudopanax arboreus and Dysoxylum spectabile) the endemic honeyeaters were most likely to meet the conditions necessary for successful pollination. For the remaining two species (Metrosideros excelsa and Geniostoma ligus trifolium) the contribution by honeyeaters and insects to pollination was equivalent. The results suggest that the role of the endemic honeyeaters in pollination of the New Zealand flora, and the subsequent regeneration of native forest ecosystems, should be important considerations in ecosystem management.     

Roles of nectar robbers in reproduction of the tropical treelet Quassia amara (Simaroubaceae)

Summary Different subsets of mainland nectarivores visited Quassia amara (Simaroubaceae), a self-compatible, predominately bird-pollinated treelet, at three islands and the mainland in Panamá. Factors correlated with reproductive success, defined as seed to ovule ratio, included the species pollinating and robbing flowers, visitor activity, pollinator response to nectar robbing, and internal regulation of fruit production. The absence of robbers and former pollinators on an island separated from the mainland during the holocene was associated with shifts in flower size, nectar production, and 3–4fold increases in population reproductive success and pollinator efficiency (=seeds produced per visit). Exclusion of robbers at three sites resulted in seed production 4–12 times greater than control flowers, at which robbers accounted for 52–98% of all visits. Although 36% of buds and over 83% of all flowers were robbed, this had no direct influence on the recorded 36–61% respective abortion rates of buds and flowers. Opportunistic avian robbers appeared where normal robbers were absent; three avian robbers extensively used floral perforations made by Trigona bees, and all ancillary pollinators also robbed. Selection pressures from nectar robbers are discussed that may relate to plant reproductive fitness.     

Temporal and microclimatic partitioning of the floral resources of Justicia aurea amongst a concourse of pollen vectors and nectar robbers

Summary The flowers of Justicia aurea, morphologically characteristic of ornithophily, attracted a diverse array of foragers where they occurred as a dense stand in the tropical forests at La Selva, Costa Rica, and so provided an arena for this study of competition and coexistence. Two hummingbird species (Phaethornis superciliosus and Campylopterus hemileucurus) visited the flowers legally early in the morning, and defended the nectar resource; a third smaller bird (P. longuemareus) foraged for nectar throughout the day but collected it illegally by piercing the corollas. In addition, nectar was harvested illegally by four species of stingless bee (Trigona) and by ants, creating a further drain on the limited floral resources.Consideration of the diurnal patterns of foraging activities in combination with a spatial axis (defined here in terms of microclimate and insolation) nevertheless showed a good separation of flower visits for the different nectarivores. Hummingbirds visited flowers in zones where the reward was highest, while insects foraged to minimise their energetic costs; each of these factors could in turn be related to microclimatic considerations. A comprehensive scheme of resource utilisation could therefore be extracted from the field observations and interpreted in these terms. The limited area of niche overlap thus revealed corresponded closely with the situations where direct interference competition was observed, between hummingbird species or between bees and aggressive ants.