Survival of the Orange-tufted Sunbird,Nectarinia osea

Cape Sugarbirds are southern African endemics, found in fynbos vegetation of the Western and Eastern Cape Provinces of South Africa, and they feed almost exclusively on the nectar of Protea inflorescences. Breeding male Sugarbirds are highly territorial and defend resources for themselves, their mates and their offspring. Sugarbirds, however, often leave their breeding territories in search of food during the dry season and return to the same breeding sites each year, thus experiencing an annual cycle of movement from one food source to another. Male territory size as well as breeding success, were determined over two breeding seasons for a population in the Helderberg Nature Reserve in the Western Cape Province. Sugarbird fledgling success was significantly greater for males defending large territories. Males moved or increased the size of their territories between seasons, possibly to improve breeding success, but no changes in territory size were observed during the breeding season.     

Farming with native bees (<Emphasis Type="Italic">Apis mellifera</Emphasis> subsp. <Emphasis Type="Italic">capensis</Emphasis> Esch.) has varied effects on nectar-feeding bird communities in South African fynbos vegetation

Outside their natural range, honeybees (Apis mellifera) are known to have detrimental effects on indigenous pollinators through exploitative or interference competition, but little is known about the effect of honeybee farming in areas where honeybees occur naturally. In the Cape Floristic Region of South Africa, where honeybees are indigenous, managed hives potentially elevate the abundance of honeybees far above natural levels, but impacts on other floral resource-dependent species have not been studied. Here we use experimental manipulation of honeybee density to test whether honeybee farming affects nectar-feeding birds. We selected the common sugarbush (Protea repens), utilized by both birds and bees, and analysed the time (before/after) by treatment (control/experiment) interaction to explore changes in bee abundance, nectar availability and bird abundance at three sites. Hive addition increased honeybee abundance in inflorescences of P. repens above expected levels. Despite experimental increase in honeybee numbers, there is no reduction in nectar sugar availability relative to the control areas. Where honeybee density was highest, sugarbird (Promerops cafer) numbers declined relative to expected, but sunbirds (Nectarinidae) were not affected at any of the sites. We conclude that stocking rates of more than one honey bee per P. repens inflorescence have detrimental effects on bird abundance due to interference, rather than resource competition.     

Sex‐related spatial patterns of Poa ligularis in relation to shrub patch occurrence in northern Patagonia

Abstract. Poa ligularis is a dioecious species and a valuable forage plant which is widespread in the arid steppe of northern Patagonia (Argentina). The vegetation in these areas consists of a system of perennial plant patches alternating with bare soil areas defining contrasting micro‐environments. We hypothesized that (1) male and female individuals of P. ligularis are spatially segregated in different micro‐environments, (2) the intensity of spatial segregation of sexes depends on plant structure and (3) spatial segregation of sexes is enhanced by competitive interactions between the sexes within the vegetation patches. We analysed the spatial distribution of female and male individuals in relation to the spatial pattern of vegetation in two areas differing in their vegetation structure. The location of P. ligularis within patches where either male, female or both sexes occurred was also analysed. The results indicate that different patterns of spatial distribution of sexes of P. ligularis may be found at the community level depending on the dominant life forms and geometric structure of plant patches. Where patches are of a lower height, with a high internal patch cover, individuals of both sexes are concentrated within patch canopies. In sites characterized by large, tall patches and less internal patch cover suitable microsites for female and male P. ligularis occur both within and outside the patch with males located at further distances from the patch edge. Where the patch is large and tall enough to allow the establishment of males and females at relatively high numbers, males occupy the patch periphery or even colonize the interpatch bare soil. These spatial patterns are consistent with selective traits in which females better tolerate intraspecific competition than males, while males tolerate wider fluctuations in the physical environment (soil moisture, nitrogen availability, wind intensity, etc.).     

3种雌雄异熟樟科植物开花动态的比较研究

为了解樟科植物雌雄异熟的繁育系统特点,对3种樟科植物阴香(Cinnamomum burmannii)、紫楠(Phoebe sheareri)和浙江楠(Phoebe chekiangensis)雌雄异熟花的开花动态进行了比较研究.结果表明,3种植物雌性功能期的开始时间、雌性功能期和雄性功能期的时间分配有差异.3种植物的主...     

Uncovering diversity of Australian Eucalyptus‐constrained felt scales (Hemiptera: Coccoidea: Eriococcidae)

Several felt scale (Eriococcidae) lineages show deep, phylogenetically constrained, host use. The most diverse of these host‐constrained lineages is Australian, associated with plants in the family Myrtaceae, and the most diverse sublineage within this group is associated with Eucalyptus. Here we use Bayesian analysis of morphological data to estimate relationships among the eucalypt‐feeding species of this group. We identify an unusual species for which we erect the monotypic genus Heathcotia Hardy & Beardsley gen.n. , with type species Heathcotia crypta Hardy & Beardsley sp.n. , and recover a clade comprised of six new species closely related to Lachnodius hirtus Maskell. We erect the genus Lobimargo Hardy & Gullan gen.n. for the latter clade, designate Lobimargo sagittisetus Hardy & Beardsley sp.n. as its type species, transfer Lachnodius hirtus to the genus as Lobimargo hirtus (Maskell) comb.n. and describe the five other new species: Lobimargo brookesae Hardy & Beardsley sp.n., Lobimargo donaldsoni Hardy & Gullan sp.n. , Lobimargo latrobeus Hardy & Beardsley sp.n. , Lobimargo rhipidotrichus Hardy & Gullan sp.n. and Lobimargo williamsi Hardy & Gullan sp.n. A lectotype is designated for L. hirtus. The adult female of each species, the first‐instar nymph of L. sagittisetus, the second‐instar female of L. hirtus and the adult male of L. williamsi are described and illustrated. A key is provided to the adult females of all eriococcid genera found on eucalypts and of all Lobimargo species. Adult females of Lobimargo have been found on stems, sometimes causing shallow pits, and on foliage or under bark, but may move to the latter habitat for oviposition.     

Patterns of plant visitation by nectar-feeding lizards

Geckos in the genus Hoplodactylus visit flowers to feed on nectar. I examined the patterns of flower visitation exhibited by two gecko species (H. maculatus and H. duvauceli) having access to two plant species: pohutukawa (Metrosideros excelsa: Myrtaceae) and flax (Phormium tenax: Agavaceae). Individual geckos were not observed to visit both plant species; individuals visiting flax tended to revisit the same plant. Geckos visiting pohutukawa were larger than those visiting flax and exhibited an early night peak in plant visitation, while lizards on flax displayed a more even pattern of activity throughout the night. On flax, geckos were more likely to be found on plants with a greater number of male flowers. Male flax flowers were of greater diameter than female flowers and produced nectar at higher rates and with greater concentrations of sugars. Experimental manipulation of pohutukawa nectar volumes suggested that the distribution of geckos is influenced by the pattern of nectar availability.     

Nectar-feeding by suburban Blue Tits: contribution to the diet in spring

The incidence of nectar feeding by 2 pairs of Blue Tits Parus caeruleus was investigated in relation to the availability of nectar and alternative food. Nectar was not the most preferred food and the occurrence of nectar feeding did not correlate significantly with most measures of nectar availability, but the Blue Tits selected the most productive flowers. Nectar feeding frequently occurred when the preferred peanuts were unavailable due to interspecific competition. The nectar resource was under-utilized by the Blue Tits except at the beginning of the flowering period. Despite this, nectar was a highly profitable food source, yielding 0.33-0.38 kJ min^?1 foraging. During the flowering period nectar was estimated to contribute up to 32.7% of the average daily metabolic rate (ADMR) of the male and up to 49.3% of the ADMR of the female, with means over the 2 years of 7.4% and 13.2% per day for the male and female, respectively. The importance of nectar in the ecology of Blue Tits is discussed.     

Nectar sugars enhance fitness in male Coenonympha pamphilus butterflies by increasing longevity or realized reproduction

The principal components of floral nectar are water and the sugars sucrose, fructose and glucose. Several studies have shown the importance of nectar sugars for female butterfly fecundity, whereas to date little attention has been paid to the effect of nectar sugars on male butterfly reproduction. Clear evidence for an effect of nectar sugars on male realized reproductive success is still missing. In this study, we fed male Coenonympha pamphilus butterflies nectar mimics with low (5%), medium (20%) or high (30%) total sugar concentrations with a sucrose:glucose:fructose ratio of 2.7:1.1:1. Sugar solutions were made mimicking Knautia arvensis, an essential nectar plant for C. pamphilus and many other European butterflies. Realized male reproductive success for each treatment was measured indirectly via nuptial gifts, by recording reproductive parameters and by characterizing time patterns over the oviposition period of their female partner. Male butterflies fed high‐concentrated nectar sugars had a longer lifespan than males fed low‐concentrated nectar sugars. In contrast, offspring of males fed medium‐concentrated nectar sugars had a higher hatching mass than progeny of males fed low‐concentrated nectar sugars, indicating a tradeoff between somatic maintenance and reproduction in the use of nectar sugars. Thus, allocation patterns of nectar sugars differed according to sugar concentrations in adult food. The method used in this experiment took into account the indispensable role of female butterflies in passing male nutrients to offspring. With this comprehensive approach, we can show the general importance of nectar sugars for male butterfly fitness and support previous findings suggesting a coevolutionary process between butterflies and flowers dependent on butterfly pollination.     

Isolation of highly polymorphic autosomal microsatellite loci and a sex‐linked locus from sugarbirds

We describe the isolation of six tetranucleotide microsatellites from the Cape sugarbird (Promerops cafer) using an enrichment protocol. All loci were highly variable with number of alleles ranging from nine to 26 and values of observed heterozygosity ranging from 0.534 to 0.931. All loci were in Hardy–Weinberg equilibrium with the exception of Pro66 and Pro86. Further analysis of Pro86 indicated it was Z‐linked. All loci amplified and were variable in the congeneric Gurneys sugarbird (Promerops gurneyi). These loci will be used in mating system and phylogeographic studies of Cape sugarbirds in South Africa.     

Pollination by deceit,floral sex ratios and seed set in dioecious Rubus chamaemorus L.

Summary Male and female flowers of the dioecious perennial herb Rubus chamaemorus L. are similar in general appearance. However, female flowers are somewhat smaller, do not produce any pollen, and contain very small amounts of nectar. Syrphids and bumblebees, which are important pollinators of R. chamaemorus, showed a strong preference for male flowers. Male flowers were also less often rejected by flower visitors than were female flowers, and two different groups of syrphid species stayed longer in male than in female flowers. These observations suggest that female flowers of R. chamaemorus attract pollinators by deceit.Hand-pollination experiments indicated that pollen availability limited seed production of R. chamaemorus in female dominated habitats but not in areas with an equal floral sex ratio. We suggest that the relative importance of factors limiting female reproductive success is not constant, but is influenced by the floral sex ratio of the population. This should apply also to other dioecious species that show variable sex ratios on either a local or regional scale.     

Can nectar properties explain sex‐specific flower preferences in the Adonis Blue butterfly Lysandra bellargus?

1. Field observations in the Swiss Jura mountains showed that males and females of the bivoltine Adonis Blue butterfly Lysandra bellargus Rott. differed significantly in their flower visitation patterns. 2. In both generations, females visited a broader range of available nectar plants than did males. The specific flower visitation patterns of males and females were not affected by the general availability and abundance of potential nectar plant species during both flight periods, indicating high selectivity for nectar plants by both males and females. 3. In addition, the sexes differed in their nectar foraging behaviours: distances between successively visited flowers were significantly longer in males than in females, indicating that male and female butterflies have different foraging strategies. 4. Investigations of nectar characteristics showed that the sexes preferred flowers with different nectar compositions. Males of both generations preferred flowers with high proportions of sucrose and high amounts of total sugar, whereas females preferred flowers with high portions of glucose in their nectar, and, in the spring generation, flowers rich in amino acids. 5. Flowers visited exclusively by males or females in spring differed significantly in their amino acid composition. 6. This clear‐cut pattern did not hold for the autumn generation, most probably due to the limited availability of flowers. 7. The observed nectar foraging patterns underline the importance of adult feeding for longevity and reproduction in butterflies. The findings are particularly relevant for conservation, because L. bellargus is an increasingly threatened species in many European countries.     

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.     

Flower choice by honey bees (Apis mellifera L.): sex-phase of flowers and preferences among nectar and pollen foragers

Bees foraging for nectar should choose different inflorescences from those foraging for both pollen and nectar, if inflorescences consist of differing proportions of male and female flowers, particularly if the sex phases of the flowers differ in nectar content as well as the occurrence of pollen. This study tested this prediction using worker honey bees (Apis mellifera L.) foraging on inflorescences of Lavandula stoechas. Female flowers contained about twice the volume of nectar of male flowers. As one would predict, bees foraging for nectar only chose inflorescences with disproportionately more female flowers: time spent on the inflorescence was correlated with the number of female flowers, but not with the number of male flowers. Inflorescence size was inversely correlated with the number of female flowers, and could be used as a morphological cue by these bees. Also as predicted, workers foraging for both pollen and nectar chose inflorescences with relatively greater numbers of both male and female flowers: time spent on these inflorescences was correlated with the number of male flowers, but not with the number of females flowers. A morphological cue inversely associated with such inflorescences is the size of the bract display. Choice of flowers within inflorescences was also influenced predictably, but preferences appeared to be based upon corolla size rather than directly on sex phase.     

The dynamics of pollen removal and deposition,and its effects on sexual phases in a protandrous plant: Glechoma longituba (Lamiaceae)

The duration of sexual phases in dichogamous plants are affected by many factors. Using both experimental and observational studies, we investigated natural patterns of pollen removal and deposition, visiting frequency of pollinators, patterns of nectar secretion, and effects of pollen removal and stigmatic pollen deposition on the duration of sexual phases in a protandrous plant, Glechoma longituba. We found that visiting frequency of pollinators correlated with the nectar secretion pattern. The nectar volume during the male phase was higher than during the female phase. In the morning, the main pollinator, the bee Anthophora plumipes, mainly foraged for nectar and showed no preference for flowers in male or female phase, despite male phase flowers producing higher amounts of nectar. However, in the afternoon, they changed their behavior and foraged mainly for pollen, and then showed a preference for flowers in male phase. Furthermore, the rates of pollen removal and stigmatic pollen deposition can affect the starting time and the duration of the female phase. When pollen removal and pollination rates are low due to scarcity of pollinator services, the sexual phase can be prolonged, leading to an overlap, and thereby enhance the chance for sexual reproduction through pollinator‐facilitated self‐pollination. We consider the variation of sexual phases in Glechoma longituba an adaptive mechanism prepared for both cross‐pollination enhancement and reproductive assurance depending on the available pollination services.     

The nectar of Justicia and Columnea: Composition and concentration in a humid tropical climate

Summary Studies in Costa Rica on two ornithophilous flowers, Justicia aurea (Acanthaceae) and Columnea glabra (Gesneriaceae) showed a constancy of nectar solute concentrations that was attributed to microclimatic protection by the tubular corolla and to copious nectar secretion, helped by waterproofing by a lipid film on the nectar surface in Justicia and by preferential compass orientation of the flowers of Columnea.Most of the corollas in the patch of Justicia had been pierced by nectar-robbers. A consequence of this damage, together with local microclimate effects, was flower-to-flower variation in the amount and accessibility of nectar and in the nature and concentration of its minor components, notably amino acids.McDade and Kinsman's (1980) finding that nectar secretion could be suppressed by repeated sampling or by nectar-robbing was confirmed.     

ADAPTIVE SIGNIFICANCE OF IPOMOPSIS AGGREGATA NECTAR PRODUCTION: OBSERVATION AND EXPERIMENT IN THE FIELD

Production of floral nectar is generally thought to be an adaptation that increases plant fitness by altering pollinator behavior, and therefore pollination success. To test this hypothesis, I investigated the effects of floral nectar production rate on pollination success of the hermaphroditic plant Ipomopsis aggregata (Polemoniaceae). Success through male function (estimated by the export of fluorescent dyes) was significantly greater for plants with naturally high nectar production rates than for nearby plants with low nectar production rates, whereas success through female function (receipt of fluorescent dye) was unrelated to nectar production rate. Experimental addition of artificial nectar also produced a significant increase in male function success and no increase in several estimates of female function success. Observations confirmed that hummingbirds probed a larger proportion of flowers on plants that received supplemental nectar, as they do in response to natural variation in nectar production. The concordance of results across these observational and experimental studies indicates that nectar production acts primarily to increase pollination success through male function for this species.     

Bumblebee-pollination and temporal change of the calyx tube length in Clematis stans(Ranunculaceae)

Clematis stans is dioecious semi-arboreal, with pale purple–blue, nodding, tubulous flowers in a paniculate inflorescence. Both male and female flowers produce nectar from the base of the calyx tube during a flowering period of 3 or 4 days, and are pollinated by two bumblebee species, Bombus diversus and B. honshuensis, with different proboscis lengths. When the flowers open, four sepals constructing a calyx tube separate at the top and their respective tips gradually curl up, so that a tubular part shortens. Observations at two field sites showed that B. diversus (with a longer proboscis) most often visits the flowers with a longer calyx tube, and B. honshuensis (with a shorter proboscis) the flowers with a shorter calyx tube, i.e., later in the flowering period. By changing the calyx tube length, the flowers of C. stans accept the two bumblebee species with different proboscis length as pollinators and thus increase the chance of pollination for each flower. It was also found that the two bumblebee species prefer the male flowers to the female flowers, although the female flowers secrete more nectar as a reward than male flowers. This is likely because they visit the male flowers to collect pollen grains in addition to nectar. Electronic Publication     

Small but attractive: female‐biased nectar production and floral visitors in a dimorphic shrub

• In sexually dimorphic species, hermaphrodite flowers in gynodioecious species or male flowers in dioecious species are often larger and produce more nectar than their conspecific female flowers. As a consequence, hermaphrodite or male flowers frequently receive more pollinator visits.
• Sex ratio, flower size, floral display, nectar production and floral visits were evaluated in two natural populations of Fuchsia thymifolia, a morphologically gynodioecious but functionally subdioecious insect‐pollinated shrub.
• Sex ratio did not differ from the expected 1:1 in the two studied populations. As expected, hermaphrodite flowers were larger than female flowers, but in contrast to the general pattern, hermaphrodite flowers did not produce nectar or produced much less than female flowers. Flower visitors were flies (68%) and bumblebees (24%), both of which showed a preference for female flowers. No sex difference was detected in either flower longevity or floral display across the flowering season.
• Higher nectar production by females may attract more pollinators, and may be a strategy to enhance female reproductive success in this species. Finally, floral dimorphism and insect preferences did not seem to hamper the maintenance of sub‐dioecy or prevent the evolution of dioecy in F. thymifolia.

     

Nectary and gender‐biased nectar production in dichogamous Chamaenerion angustifolium (L.) Scop. (Onagraceae)

In dichogamous plants, nectar characteristics (i.e. nectar amount and its composition) can differ between sexual phases. In the present study, we investigated the structural organization of the floral nectary, nectar production and carbohydrate composition in the protandrous Chamaenerion angustifolium (L.) Scop. (Onagraceae). The receptacular nectary consisted of an epidermis with numerous nectarostomata, several layers of photosynthetic secretory parenchyma, and subsecretory parenchyma. Nectariferous tissue was not directly vascularized and starch grains were rarely observed in the secretory cells, occurring exclusively in the guard cells of modified stomata. The nectar was released via nectarostomata. The floral nectar was hexose rich (32.8/39.1/28.1% glucose/fructose/sucrose) and the total concentration was constant throughout the anthesis (47% on average). However, contrasting patterns in nectar amount and carbohydrate composition between the floral sexual phases were observed. On average, female‐phased flowers produced 1.4‐fold more nectar than male‐phased flowers, and although the nectar was sucrose rich during the male phase, it was hexose rich during the female phase, suggesting sucrose hydrolysis.     

OCCURRENCE AND MOVEMENT OF MIGRATORY SPECIES IN RHODESIA AND SOUTHERN AFRICA DURING THE PERIOD 1950-1953

Summary

Seiler, H. W. &; Pr?s-Jones, R. P. 1989. Mate competition, mate guarding, and unusual timing of copulations in the Cape Sugarbird (Promerops cafer). Ostrich 60:159-164.

Focal observations on two breeding pairs of the Cape Sugarbird were undertaken in the Cape Peninsula, South Africa. During the presumed fertile period of the female the male was significantly less likely to become separated from his mate than during other stages of the breeding cycle. This is interpreted as mate paring, which might be triggered by the female well in advance of her actual fertile period. The presumed fertile period was closely accompanied by a sharp increase in territorial intrusions by other males. Copulations were observed only outside the presumed fertile period.