Competition for nectar resources does not affect bee foraging tactic constancy

1. Competition alters animal foraging, including promoting the use of alternative resources. It may also impact how animals feed when they are able to handle the same food with more than one tactic. Competition likely impacts both consumers and their resources through its effects on food handling, but this topic has received little attention. 2. Bees often use two tactics for extracting nectar from flowers: they can visit at the flower opening, or rob nectar from holes at the base of flowers. Exploitative competition for nectar is thought to promote nectar robbing. If so, higher competition among floral visitors should reduce constancy to a single foraging tactic as foragers will seek food using all possible tactics. To test this prediction, field observations and two experiments involving bumble bees visiting three montane Colorado plant species (Mertensia ciliata, Linaria vulgaris, Corydalis caseana) were used under various levels of inter- and intra-specific competition for nectar. 3. In general, individual bumble bees remained constant to a single foraging tactic, independent of competition levels. However, bees that visited M. ciliata in field observations decreased their constancy and increased nectar robbing rates as visitation rates by co-visitors increased. 4. While tactic constancy was high overall regardless of competition intensity, this study highlights some intriguing instances in which competition and tactic constancy may be linked. Further studies investigating the cognitive underpinnings of tactic constancy should provide insight on the ways in which animals use alternative foraging tactics to exploit resources.     

Flower change in Oenothera drummondii Hooker as a response to pollinators' visits

Flower change in Oenothera drummondii Hooker as a response to pollinators' visits. The colour and shape of flowers of Oenothera drummondii change owing to senescence, pollination and/or nectar withdrawal. This phenomenon is interpreted as a signal emitted by the plant to 'inform' the potential pollinator of its new status as a rewardless flower. This method of signalling might be important for the pollinators in order to save energy in seeking the correct flowers and for the plant to ensure visits to those flowers that have not yet been visited.     

Taxonomic studies ofAsarum sensu lato

The floral vascular anatomy of 12 species representing each ofAsarum s. str.,Asiasarum, Geotaenium, Heterotropa andHexastylis are compared to clarify intergeneric relationships. The five genera have basically similar structures in floral morphology and vasculature, and consistently have a six-carpelled compound ovary and the associated similar placental vasculature. They show, however, a significant difference in the position and the constituent of the “ventral” carpellary bundles in the placental axis betweenAsiasarum-Heterotropa-Hexastylis andAsarum-Geotaenium. InAsiasarum, Heterotropa andHexastylis the ventral bundles of each carpel are basically free and antilocular as expected in the least specialized compound ovary of angiosperms; in contrast, inAsarum andGeotaenium the ventral carpellary bundles are antiseptal and heterogenous (i.e., formed by the lateral fusion of ventral bundles of adjacent carpels). Shared probable apomorphic floral vasculature, as well as shared single style-column, suggests the closest mutual relationships betweenAsarum andGeotaenium. In terms of floral morphology and anatomy,Asiasarum, Heterotropa andHexastylis retain plesiomorphies. Possible chromosomal evolution in the related genera is also discussed.     

Determinants of foraging profitability in two nectarivorous butterflies

ABSTRACT.

• 1 I studied flower selection and foraging energetics of Agraulis vanillae L. (Nymphalidae) and Phoebis sennae (Pieridae), two butterfly species common to north central Florida. I identified the major nectar resources exploited by several populations of these butterflies and, for each plant species, measured available nectar volumes and concentrations, corolla lengths, and density. I quantified foraging behaviour of each butterfly species at each nectar source (flower visitation rate and percentage of foraging time in flight), and used these data to estimate the net rate of energy intake of each butterfly species at each nectar source.
• 2 Estimated mean energy contents of individual flowers of the eleven exploited plant species spanned three orders of magnitude, ranging between 0.015 and 9.27 joules. Mean energy content of individual flowers was strongly correlated with mean foraging profit of both butterfly species.
• 3 Mean nectar volume strongly influenced energy content and varied widely within and among species, ranging from 0.0076 to 1.853 μ1. Nectar concentration varied between 17.1% and 40.4% sucrose-equivalents. Nectar volume was the best single predictor of foraging profitability (correlation coefficients of 0.994 and 0.984 for Phoebis and Agraulis respectively). Corolla length also strongly affected foraging profitability for both butterfly species; flower species with longer corollas were generally more profitable.
• 4 Flower density and nectar concentration showed weak or nonsignificant associations with foraging profitability.
• 5 The usefulness and limitations of these floral characteristics as bases for foraging selectivity, and the selective pressures foraging butterflies might place on the visited plants are discussed.

     

Estimates of hierarchical variation in flower morphology in natural populations ofScleranthus annuus (Caryophyllaceae), an inbreeding annual

The variation in two sets of morphological characters of the flowers of the highly inbreedingScleranthus annuus (Caryophyllaceae) was assessed using 15–20 plants from each of 20 natural populations from the southernmost region of Sweden. The stamen fertility data set consisted of 10 characters describing the degree of fertility of the ten stamens/staminoids, while the sepal/gynoecium data set comprised 10 sepal characters, together with style and stigma length. Substantial variation was found in both the degree of development and the fertility of the stamens, the degree of variability in stamen fertility being related to stamen position within the flower. Considerable variation was found in the characters of the sepal and the gynoecium. Hierarchical analyses of variance indicated that 29% of the variation in total male reproductive effort was distributed among populations, 28% among plants within populations and 43% represented within-plant variation. The corresponding averages for the characters from the sepal/gynoecium data set are 26, 38, and 35%: a greater proportion of the total variance in female reproductive characters is accounted for by among individual variation than is the case with the male reproductive characters. Significance tests of Mahalanobis distances derived by canonical variate analyses indicated that all populations were significantly separated using the sepal/gynoecium data set, while only 50% of the pairwise comparisons on the basis of the stamen fertility data set were significant. Cluster analysis did not reveal any aggregation of the populations. The incongruence of the two data sets and their ability to discriminate between the populations is discussed.     

Reproductive structures and phylogenetic significance of extant primitive Angiosperms

A brief survey is presented on fossil reproductive structures of early Angiosperms from the Lower and mid-Cretaceous and at the same time on the reproductive structures of those extant Angiosperms which resemble most closely these fossils and which seem to be especially primitive also on other grounds: a first group (Degeneriaceae, Himantandraceae, Eupomatiaceae, Austrobaileyaceae) possessing relatively complicated and conspicuous flowers with elaborated inner staminodes, a second group (Chloranthaceae, Trimeniaceae, Amborellaceae) possessing small and relatively simple, inconspicuous flowers with peculiar features in the carpels, and a third group (Winteraceae) possessing flowers with unusual variability in organ number and size. The three groups exhibit a certain diversity in pollination biology, although cantharophily seems to prevail, however different the cantharophily character syndromes may be between the groups. In the extant primitiveMagnoliidae variability occurs on other morphological levels than in the higher advanced Angiosperms. This has to be taken into consideration in evaluations of the systematic relationships of the various groups of theMagnoliidae. Presumably often their relationships are closer than it may appear at first sight. This is also true for the three groups here discussed.     

Studies in the floral morphology and anatomy of nymphaeales

Floral morphology ofBrasenia schreberi Gmel. andCabomba caroliniana A. Gray was observed chiefly from an anatomical point of view. The receptacle ofB. schreberi is rather flat and a vascular plexus is observable in the mature flower. The vasculature in this plexus is so complex taht it is not easy to trace its structure in detail. by observation on small buds, it can be seen that the receptacular vasculature consists of a girdling bundle in the basal area and usually nine receptacular strands from which traces to the petals and stamens branch off. The vasculature in the receptacle is reconstructed and diagramatically shown as though split longitudinally and spread out in one plane. Floral vasculature inCabomba caroliniana is simpler, and is probably related to the smaller number of stamens and carpels. It also has a girdling bundle at the bottom of receptacle and this vasculature is suggested to be derived by simplification from aBrasenia-type vasculature. Evidence from floral anatomy suggests that these two genera are closely related. InNymphaea, a vascular plexus in the receptacle is also observed (Moseley, 1961; Ito 1983). The plexus ofBrasenia andNymphaea are not the same in their construction. Nevertheless, their fundamental floral vasculature is comparable and it is preferable to place them in the same family or same order.     

Studies in Garcinia, dioecious tropical forest trees: the phenology, pollination biology and fertilization of G. hombroniana Pierre

RICHARDS, A. J., 1990. Studies in Garcinia , dioecious tropical forest trees: the phenology, pollination biology and fertilization of G. hombroniana Pierre . Garcinia hombroniana is a facultative agamosperm which is pollinated by Trigona bees. Nectar is restricted to the large discoid stigma (or pistillode in male flowers), which also captures and hydrates pollen. The 'wet' stigma and binucleate pollen suggest that Garcinia arose from hermaphrodite plants with a gametophytic self-incompatibility system.
On stigmas, nectar is secreted early on three or four successive days. On male pistillodes, nectar is secreted when anthers dehisce, on the second morning after anthesis. Pollen is most viable when freshly collected, but some viability remains four days after collection. Pollen germinates within 24 h of hydration. Similar results to pollinations are obtained by germinating pollen in 1 % sucrose.
Garcinia hombroniana flowers principally from January to June. Cultivated females are considered as 'big bang' strategists. Male flowers are considered as 'steady state' strategists.     

EVOLUTION OF FLORAL TRAITS IN A HERMAPHRODITIC PLANT: FIELD MEASUREMENTS OF HERITABILITIES AND GENETIC CORRELATIONS

Genetic variances, heritabilities, and genetic correlations of floral traits were measured in the monocarpic perennial Ipomopsis aggregata (Polemoniaceae). A paternal half-sib design was employed to generate seeds in each of four years, and seeds were planted back in the field near the parental site. The progeny were followed for up to eight years to estimate quantitative genetic parameters subject to natural levels of environmental variation over the entire life cycle. Narrow-sense heritabilities of 0.2–0.8 were detected for the morphometric traits of corolla length, corolla width, stigma position, and anther position. The proportion of time spent by the protandrous flowers in the pistillate phase (“proportion pistillate”) also exhibited detectable heritability of near 0.3. In contrast, heritability estimates for nectar reward traits were low and not significantly different from zero, due to high environmental variance between and within flowering years. The estimates of genetic parameters were combined with phenotypic selection gradients to predict evolutionary responses to selection mediated by the hummingbird pollinators. One trait, corolla width, showed the potential for a rapid response to ongoing selection through male function, as it experienced both direct selection, by influencing pollen export, and relatively high heritability. Predicted responses were lower for proportion pistillate and corolla length, even though these traits also experienced direct selection. Stigma position was expected to respond positively to indirect selection of proportion pistillate but negatively to selection of corolla length, with the net effect sensitive to variation in the selection estimates. Anther position also was not directly selected but could respond to indirect selection of genetically correlated traits.