Components of pollination effectiveness in Psychotria suerrensis,a tropical distylous shrub

In this paper I report components of effectiveness for pollinators of a tropical distylous shrub, Psychotria suerrensis (Rubiaceae), which is visited by a variety of bees, wasps, and butterflies, and by two species of hummingbirds. In the field, I measured the following components of effectiveness: frequency of visits, evenness of visits across plants, and diurnal pattern of visits. I also used flight-cage experiments to compare pollentransfer abilities of euglossine bees and heliconiid butterflies. Euglossine bees visited more frequently, visited earlier in the day, and visited a higher proportion of plants in the population than did other taxa. In flight cage experiments, bees and butterflies transferred similar amounts of pollen overall, but bees transferred significantly more inter-morph (compatible) pollen. For each component measured, euglossine bees appeared to be the most effective pollinators.     

Seed set in Phormium: interactive effects of pollinator behaviour,pollen carryover and pollen source

Summary Fruit and seed set was studied in New Zealand flax (Phormium tenax Phormiaceae) a large monocot that preferentially sets outcrossed seeds. Fruit set was low and in particular situations could result from insufficient pollinator visitation. Observations of pollinator (Meliphagidae) movements showed that birds preferentially visited male phase flowers and predominantly moved pollen within inflorescences of the same plant. More dominant resident birds moved more between plants that subordinate non-resident birds. Combination of results of fluorescent dye carryover with known bird movements allowed predictions of fruit set and seed size that closely approximated observed levels. Resident birds account for almost half the observed foraging bouts but are predicted to be responsible for the vast majority of the viable seeds.     

Electroantennogram responses of the Mediterranean fruit fly, Ceratitis capitata to identified volatile constituents from calling males

Fifty-six compounds from the odor of calling, sexually mature, laboratory reared males of the Mediterranean fruit fly, Ceratitis capitata (Wiedemann) (Diptera: Tephritidae) were isolated by headspace trapping on Tenax columns and identified using GC/MS techniques (69 total compounds were detected). Electroantennogram responses (EAGs) to 54 of the 56 identified compounds as well as 5 analogs were tested on both sexes. Significant differences between the sexes in their responsiveness were found in 9 of the 54 identified compounds tested. There was no correlation between the amplitude of the EAG response and the relative abundance of compound identified from headspace analysis. Of the five major identified components, three elicited relatively small EAG responses, while two elicited large EAGs compared to the hexan-1-ol standard. The relative ranking of EAG responses were: methyl and ethyl hexenoates and hexanoates > C₄–C₆ esters and/or acetates > ethyl and methyl octenoates > monoterpenes > sesquiterpenes > C₂–C₅ acetates, alcohols and ketones. Behavioral bioassays on each of the five major identified components as well as a blend of six of the compounds showed some degree of attractancy to virgin females which in some cases approached the response to a pheromonal standard (male odors absorbed onto filter paper). These results are discussed in relationship to the insect's antennal sensitivity to putative pheromone components and/or allomonal components and to other reported C. capitata pheromone studies.

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Résumé Cinquante-six composés de l'odeur de mâles de C. capitata Weidemann, élevés en laboratoire, sexuellement mûrs et en appel, ont été isolés par piégeage sur colonnes tenax et identifiés par la technique GC/MS (69 composés avaient été détectés en tout). Les électroantennogrammes (EAGs) ont été examinés chez les deux sexes pour 54 des 56 composés identifiés et 5 de leurs analogues. Des différences significatives entre les sexes ont été observées pour 9 des 54 composés identifiés. Il n'y avait pas de corrélation entre l'ampleur de l'EAG et l'abondance relative du composé lors de son isolement. Pour les 5 principaux composés identifiés, 3 ont induit des EAGs relativement faibles, tandis que 2 étaient importants, par comparaison avec l'Hexane-1-ol utilisé comme témoin. Le classement relatif des EAG a été: hexénoates et hexanoates d'éthyl et de méthyl C₄–C₆ esters et/ou acétates octénoates d'éthyl ou de méthyl monoterpènes sesquiterpènes C₂–C₅ acétates, alcools et kétones. Les expériences de comportement avec chacun des 5 composés principaux identifiés, comme avec des mélanges de 6 composés ont mis en évidence une attraction des femelles vierges qui dans quelques cas avoisine la réponse à la phéromone témoin (odeur du mâle absorbée sur papier filtre). Ces résultats sont discutés en fonction de la sensibilité de l'antenne d'insexte aux composés supposés de la phéromone et aux composés allomonaux, et en fonction des autres études connues sur les phéromones de C. capitata.

     

Stamens are not essential as an attractant for pollinators in females of cryptically dioecious Thalictrum pubescens Pursch. (Ranunculaceae)

 It has been hypothesized that females of some dioecious species maintain stamens that produce sterile pollen as a means of attracting pollinators to promote greater seed set. However, this hypothesis has rarely been tested. This paper examines the role of pollinators in the maintenance of sterile stamens in the cryptically dioecious species, Thalictrum pubescens. Wind was found to contribute to pollination; branches of female T. pubescens enclosed in cheesecloth to exclude insects but not wind were still able to set seed. Therefore, females may not need the stamens for pollination. In 1994, insects were found to discriminate against emasculated female flowers, but this discrimination did not lead to a significant difference in either the amount of pollen received or seed set. In 1995, emasculation was combined with the addition of pollen to determine if emasculated females were pollen limited. No significant effect of pollen addition or emasculation was found. Emasculated branches in the pollen-addition treatment appeared to have slightly lower seed set than non-emasculated branches that received pollen, indicating that the slight reduction in seed set was caused by emasculation rather than pollen limitation. Since stamens do not appear to be maintained as pollinator attractants, other explanations, such as genetic constraints, must be investigated. Received: 4 February 1997 / Revision accepted: 20 May 1997     

Gene flow inferred from seed dispersal and pollinator behaviour compared to DNA analysis of restriction site variation in a patchy population of Lotus corniculatus L.

Summary Gene flow was investigated in a natural population of Lotus corniculatus L. (Fabaceae) using a combination of pollen and seed dispersal studies and a recombinant DNA technique. The population is spatially heterogeneous and grows with Empetrum nigrum. L. corniculatus is pollinated by the pollen-collecting bumblebee Bombus lapidarius L. Most pollinator flights occurred within patches, as bees usually visit nearest-neighbour plants, show no marked directionality, and forage mostly within patches. Gene flow by seeds is also limited, reinforcing the pattern of gene flow within patches. However, 2.6% of pollinator flights are between patches and considerable pollen carryover also occurs. Thus, gene flow between patches is potentially sufficient to retard or prevent genetic differentiation in spite of the patchy sub-structuring of the population. A sub-set of the population was analysed for restriction fragment length polymorphisms (RFLPs) to document the actual gene flow pattern of the population. The DNA analysis revealed significant levels of genetic differentiation between the patches. The level of gene flow that can be inferred from the distribution of genetic variation is surprisingly restricted, as compared to gene flow inferred from pollinator behaviour, and emphasizes that stochastic processes like genetic drift and founder effects may have a strong impact on the prevailing genetic structure.     

Attraction of the grasshopper, Melanoplus sanguinipes, to host plant odors and volatile components

The attraction of the polyphagous grasshopper, Melanoplus sanguinipes (F.), to odors from plant foliage or to chemical components of these odors was tested in a glass Y-tube olfactometer. Humidified air was passed at equal flow rates through a sample chamber containing plant material and an empty control chamber and then through the Y-tube to the holding chamber containing the test insect. Solutions of volatile chemicals in water were metered at a constant rate into the sample air stream with a syringe pump. Insects moving upwind were recorded after entering either the sample or control arm of the Y-tube. Both nymphs and adults were strongly attracted to the odor of chopped and intact seedling foliage of perennial ryegrass and wheat. Chopped leaves but not intact leaves of sorghum and alfalfa also were significantly attractive. The major components of ryegrass odor from both chopped and stem-cut leaves were Z-hex-3-en-1-yl acetate, Z-hex-3-en-1-ol, and pent-1-en-3-ol, in that order, with lesser amounts of E-hex-2-enal. Chopped leaves released many more minor components. The major components when tested individually or in binary and ternary mixtures were significantly attractive to grasshoppers compared to humidified air but not to the degree of whole plant odor. However, a quartenary mixture simulating the odor blend had levels of attractancy equal to that of chopped grass odor. Blends of these 5- and 6-carbon unsaturated alcohols, esters, and aldehydes volatilizing from green plants probably play an important role as olfactory cues for orientation of grasshoppers to food plants.

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Résumé L'attraction de M. sanguinipes (F.) par l'odeur du feuillage et des composés chimiques de ces odeurs a été examinée dans un olfactomètre en tube de verre en Y. Des flux identiques d'air humidifié étaient introduits dans une enceinte contenant le matériel végétal et dans une enceinte témoin, et, via le tube en Y, dans l'enceinte contenant l'insecte à étudier. Des solutions aqueuses de substances volatiles étaient introduites à un taux constant dans le courant d'air grâce à une seringue. Les insectes se déplaçant contre le vent étaient enregistrés après pénétration dans une branche déterminée du tube en Y. Larves et adultes étaient fortement attirés par l'odeur de feuillage intact ou coupé de semis de Lolium perenne et de blé (Triticum aestivum). Les feuilles coupées, mais non les feuilles intactes de sorgho (Sorghum bicolor) et de luzerne (Medicago sativa) étaient, elles aussi, significativement attractives. Les principales substances constituant de l'odeur de feuilles de Lolium perenne, tant coupées qu'intactes, étaient dans l'ordre de Z-hex-3-en-1-yl acétate, le Z-hex-3-en-1-ol et le pent-1-en-3-ol, et dans une moindre mesure, E-hex-2-enal. Les feuilles coupées libéraient beaucoup plus de composés secondaires. Les principaux constituants testés, seuls ou en mélanges binaires ou ternaires, étaient significativement attractifs, mais moins que l'odeur totale de la plante. Toutefois, un mélange quaternaire, simulant le mélange odorant, avait le même pouvoir attractif que l'odeur de la plante entière. L'union de ces alcools, esters et aldéhydes à 5- et 6-carbones non saturés, volatilisés à partir des plantes vertes, jouent probablement un grand rôle comme signaux orientateurs lors de l'attraction des criquets par leur plantes alimentaires.

     

Flowering mechanisms,pollination strategies and floral scent analyses of syntopically co‐flowering Homalomena spp. (Araceae) on Borneo

In this study, the flowering mechanisms and pollination strategies of seven species of the highly diverse genus Homalomena (Araceae) were investigated in native populations of West Sarawak, Borneo. The floral scent compositions were also recorded for six of these species. The selected taxa belong to three out of four complexes of the section Cyrtocladon (Hanneae, Giamensis and Borneensis). The species belonging to the Hanneae complex exhibited longer anthesis (53–62 h) than those of the Giamensis and Borneensis complexes (ca. 30 h). Species belonging to the Hanneae complex underwent two floral scent emission events in consecutive days, during the pistillate and staminate phases of anthesis. In species belonging to the Giamensis and Borneensis complexes, floral scent emission was only evident to the human nose during the pistillate phase. A total of 33 volatile organic compounds (VOCs) were detected in floral scent analyses of species belonging to the Hanneae complex, whereas 26 VOCs were found in samples of those belonging to the Giamensis complex. The floral scent blends contained uncommon compounds in high concentration, which could ensure pollinator discrimination. Our observations indicate that scarab beetles (Parastasia gestroi and P. nigripennis; Scarabaeidae, Rutelinae) are the pollinators of the investigated species of Homalomena, with Chaloenus schawalleri (Chrysomelidae, Galeuricinae) acting as a secondary pollinator. The pollinators utilise the inflorescence for food, mating opportunities and safe mating arena as rewards. Flower‐breeding flies (Colocasiomyia nigricauda and C. aff. heterodonta; Diptera, Drosophilidae) and terrestrial hydrophilid beetles (Cycreon sp.; Coleoptera, Hydrophilidae) were also frequently recovered from inflorescences belonging to all studied species (except H. velutipedunculata), but they probably do not act as efficient pollinators. Future studies should investigate the post‐mating isolating barriers among syntopically co‐flowering Homalomena sharing the same visiting insects.     

The effect of feeding status on sexual attractiveness of Ixodes ricinus (Acari: Ixodidae) females

Sexual attractiveness of unengorged, semiengorged and fully engorged females of both field and laboratory populations of Ixodes ricinus was studied under laboratory conditions by means of a computerised video tracking system. A male and a female were allowed to walk freely in a glass arena during 1 h and their behavioural interactions were observed. Obtained results revealed that the feeding status of I. ricinus females affects their sexual attractiveness. The highest attractiveness was observed in engorged females, the lowest in unengorged females of the field population. Copulation occurred in all experimental groups with the highest frequency observed in the unengorged virgin laboratory females. Despite engorged females being highly attractive for males on distance, the frequency of copulation was low. Differences between field and laboratory populations were found in both behavioural parameters and frequency of copulation. This revised version was published online in July 2006 with corrections to the Cover Date.     

A test of the effect of floral color change on pollination effectiveness using artificial inflorescences visited by bumblebees

Floral color change has been recognized as a pollination strategy, but its relative effectiveness has been evaluated insufficiently with respect to other floral traits. In this study, effects of floral color change on the visitation pattern of bumblebees were empirically assessed using artificial flowers. Four inflorescence types were postulated as strategies of flowering behavior: type 1 has no retention of old flowers, resulting in a small display size; type 2 retains old flowers without nectar production; type 3 retains old flowers with nectar; and type 4 retains color-changed old flowers without nectar. Effects of these treatments varied depending on both the total display size (single versus multiple inflorescences) and the pattern of flower-opening. In the single inflorescence experiment, a large floral display due to the retention of old flowers (types 2–4) enhanced pollinator attraction, and the number of flower visits per stay decreased with color change (type 4), suggesting a decrease in geitonogamous pollination. Type-4 plants also reduced the foraging time of bees in comparison with type-2 plants. In the multiple inflorescence experiment, the retention of old flowers did not contribute to pollinator attraction. When flowering occurred sequentially within inflorescences, type-4 plants successfully decreased the number of visits and the foraging time in comparison with type-2 plants. In contrast, floral color change did not influence the number of visits, and it extended the foraging time when flowering occurred simultaneously within inflorescences but the opening of inflorescences progressed sequentially within a plant. Therefore, the effectiveness of floral color change is highly susceptible to the display size and flowering pattern within plants, and this may limit the versatility of the color change strategy in nature.