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0 - Maaike de Vias MSc thesis March 2001-January 2002 Supervision: Manja Kwak & Frank Hoff mann Laboratory of Plant Ecology Rijksuniversiteit Groningen Voorwoord Aan het eind van 2000 was mijn specialisatie fase in biologie (subtase 2) bijna afgerond. 1k moest als onderdeel voor subfase 3 een onderzoeksonderwerp plannen. Dit eerste onderzoeksonderwerp moest 22 weken duren en intern zijn. 1k wilde graag een plantenecologisch onderzoek doen. Mijn studieleider was Jelte van Andel, professor plantenoecologie aan de Rijksuniversiteit Groningen. 1k vertelde hem, wat 1kgraagzou willen: geen laboratoriumwerk, veel veidwerk en een beetje experimenteel werk in de kas. Verder wilde 1k vrij zijn in de onderzoeksvragen. 1k wilde mijn eigen onderzoek doen, met een begin en eind. Jelte stelde voor diezelfde middag naar praatjes te luisteren. Frank Hoffmann, een AlO die met Manja Kwak aan bestuivingsoecologie onderzoek deed, hield een verhaal over bestuiving door insecten in wegbermen. Mijn interesse was gewekt, en ik begon in maart met mijn onderzoek. Tot eind april bloeide er nauwelijks lets. 1k heb veel gelezen en een onderzoeksvoorstel geschreven. Hondsdraf, een Iipbloemige, was één van de bloeiende plantensoorten in april. 1k wilde de resultaten gaan vergelijken met resultaten van een andere Iipbloemige, Wife Dovenetel. Verder hadden we het idee om Zevenblad en Fluitenkruid (schermbloemigen) te onderzoeken. Deze soorten zouden een totaal verschillende insectenbezoekersgroep hebben dan de Iipbloemigen. Uiteindelijk heb 1k Hondsdraf uitvoerig en Fluitenkruid minder uitvoerig onderzocht. Daarbij gebruikte ik bermen die Manja en Frank ook gebruikten. 1k heb een paar extra proeven gedaan met Witte Dovenetel en Zevenblad. Toen het veldwerk klaar was, moest 1k flog stuifmeelkorrels in een enorme stapel preparaten tellen. Op 1 oktober heb 1k een deal van mijn resultaten gepresenteerd. Dit was de officiOle einddatum van mijn onderzoek. Het versiag bleek nog lang niet at te zijn. Andere projecten drongen zich aan. Nu, begin februari 2002, is het verslag klaar. Achteraf is het een waardevol bezit. Toch ben 1k blij, dat 1k straks weer met een schone lel kan beginnen aan de rest van mijn studie. Als alles gaat volgens planning, studeer 1k in december at, lets meer dan vijf jaar na het begin van mijn studie. Maaike, Haren, 7 februari 2002
Front page: Glechoma hederacea (left) and Anthriscus sylvestris (right) with some of their visitors. The picture is not on scale. The visitors around G. hederacea are from left to right: Apis mellifera (honey bee), Melanostoma spiPlatycheirus sp. (small hovertty), Rhingia campestris (larger hoverfly) and Bombus pascuorum (a bumblebee). The insects around Anthriscus sylvestris are: Eristalis tenax/pertinax (hovertly), Muscasp.(housefly), Helophilus sp. (hoverfly) and Scatophaga stercoraria (dung fly). Not all visitors of the plant species are shown. The arrows indicate the possibility of different effectiveness of insect species for pollination and are fictive. The edge is filled with on the left mostly G. hederacea pollen grains, and on the right with mostly A. sylvestris pollen grains. Also other pollen occur in the edge.
In this report, scientific names have been used wherever possible for plant and insect species. Some insects could not well be identified, in that case the family or order name is given. 2
Contents Chapter Page Voorwoord 1 Contents 2 Key-words 3 Abstract 3 Samenvatting 4 1. Introduction 5 2. Materials and methods 7 2.1 Background information 7 2.2 Visitors 7 2.3 Effectiveness of visitors 9 3. Results 11 3.1 Glechoma hederacea Background information 11 Visitors 18 Effectiveness of visitors 22 3.2 Anthrlscus sylvestris Background information 34 Visitors 36 Effectiveness of visitors 39 4. Conclusion and discussion Specialisation levels 47 Glechoma hederacea 48 Anthriscus sylvestris 54 General 56 5. Acknowledgements 58 6. References 58 Appendix A. Research calendar Appendix B. Verge visitor composition 2000 Appendix C. Nectar production by Glechoma hederacea plant types Appendix D. Pollen grains on Glechoma hederacea virgin pistils Appendix E. Glechoma hederacea breeding system experimental Appendix F. Breeding system Glechoma hederacea field Appendix G. Anthriscus sylvestris breeding system experimental Appendix H. Nectar Glechoma hederacea Appendix I. Nectar Lamium album Appendix J. Plot observations (10 mm) on Glechoma hederacea Appendix K. Plot observations (10 mm) on Ant hriscus sylvestris Appendix L. Transect observations Glechoma hederacea Appendix M. Transect observations Anthriscus sylvestris Appendix N. Following observations on Glechoma hederacea Appendix 0. Following observations on Anthriscus sylvestris Appendix P. Pollen load slides Glechoma hederacea Appendix Q. Pollen load slides Anthriscus sy!vestns Appendix R. Slides of pollen deposition on virgin Glechoma hederacea pistils Appendix S. Fluorescent dye powder Glechoma hederacea Appendix T. Fluorescent dye powder Anthriscus sylvestns Appendix U. Discrimination experiment with fluorescent dye powder on Aegopodium podagraria Appendix V. Weather conditions during observation days (estimations) Appendix W. List of figures and tables Appendix X. Glechoma hederacea Appendix V. Ant hnscus sylvestris Appendix Z. Thesis description 3
Key-words Glechoma hederacea, Anthriscus sylvestris, visitor guild, insect pollination,pollinator effectiveness, fluorescent dye powder, pollen load, pollen deposition Abstract Glechoma hederacea (Lamiaceae) and Anthriscus sylvestris (Umbelliferae) are commonspecies in the Netherlands. Both species canbepollinated by insects. G. hederacea has zygomorphic flowers. Nectar is easily accessible. The visiting group of insect species is called the visitorguild. Each insect species has his own pollination effectiveness. This effectiveness depends on(behavioural) characteristics: pollen load, pollen deposition, foraging speed, and flight distance. The composition ofthe visitor guild and the frequency of the visits of the vanous insect species combined with the effectivenessleads to the relative importance of visitors. Research was done in road verges in the surroundings of Assen in the Netherlands.There were three areas with differing agricultural activity. The visitor guilds wereinvestigated by plot observations and transect observations. Glechoma hederacea was split into plant types withfemale or hermaphroditic flowers. Both G. hederacea and A. sylvestns tumed out to be self-compatible by a breedingsystem experiment. Glechoma hederacea produced nectar faster during the day than during thenight. Lamium album, another common species, produces four times as much nectar, andis thus more attractive for pollinators. The most common visitor on G. hederacea (both plant types) was Apis melifera.Rhingia campestns was most abundant species on hermaphroditic flowers duringtransect walks. The number of visits per plant did not differ in three areas with increasing agricultural activity.Extra hand pollination in one verge showed that at least two visits are necessary for maximumseed set in that area. Effectiveness was mostly investigated by following individual flower visiting insects. For each trait,different insect species had the most favourable value. A fluorescent dye powder (= pollen analogue)experiment showed, that pollen grains can be transported over 64 meters in two days. For A. sylvestris, the same methods were used. The main visitors (walking transectobservations) were Enstalis tenax/pertinaxandEmpis tessellata. Plot observations said that E. tessellata was the main visitor. For each trait, other insect species had the most favourable values.Fluorescent dye powder showed that pollen grains could be transferred over 100 meters in a day. Overall, insectsseemed not to have preferences for fluorescent coloured umbels. On another common umbellifer,Aegopodium podagraria, the main visitor was Helophilus sp.. Anthnscus sylvestns and hermaphroditicG. hederacea had halfspecialised, and female G. hederacea had mostly specialised visitors. Results show that each insect species has its own contribution to cross-pollination: either it deposits a lot of conspecific pollen or flies large distances between two consecutivevisits, etc. When relative effectiveness is used and combined with relative abundance of the visitors,Syrphidae rest and Bombus pascuorum were the most important pollinators for female G.hederacea. Rhingia campestris was the most important pollinator for hermaphroditic G. hederacea. Eristalis tenaxipertinax wasthe most important pollinator for A. sy!vestns. 4
Samenvatti ng Glechoma hederacea (Hondsdraf, Lamiaceae) en Anthriscussylvestris(Fluitenkruid, Umbelliferae) zijn algemene soorten in Nederland. Ze kunnen door insecten worden bestoven. G. hederacea heeft zygomorfe bloemen. Nectar wordt gepresenteerd in een kroonbuis van ongeveer vijf millimeter diepte. Anthriscus sylvestris heeft actinomorfe bloemen en nectar zit ondiep. De bezoekende groep insectensoorten wordt het bezoekersgilde genoemd. Elke insectensoort heeft haar eigen bestuivingeffectiviteit. Deze effectiviteit hangt at van (gedrag-)kenmerken van insecten: de stuifmeel lading, stuifmeel depositie, foerageersnelheid en vliegafstanden. De samenstelling van het bezoekersgilde en de frequentie van de bezoeken van de verschillende insectensoorten gecombineerd met effectMteit leidt tot de relatieve belangrijkheid van bezoekers. Onderzoek werd in bermen rond Assen gedaan. Er waren dne gebieden met verschillende mate van landbouwactiviteit. De bezoekersgilden werden onderzocht door plotwaamemingen en transect waamemingen. Glechoma hederacea werd opgesplitst in planten met vrouwelijke of met hermafrodiete bloemen. Zowel G. hederacea als A. sylvestris bleken bij een voortplantingssysteem experiment zeltcompatibel te zijn. Glechoma hederacea produceerde overdag sneller nectar dan 's nachts. Lamium album, een andere algemene soort, produceerde vier keer zoveel nectar, en is daardoor aantrekkelijker voor bestuivers. De meest algemene bezoeker van G. hederacea was de honing bij Apis mellifera. Rhingia campestris was de meest voorkomende soort op hermafrodiete bloemen (transect waarnemingen). Het aantal bezoeken per plant verschilde niet in drie gebieden met toenemende landbouwactiviteit. Extra handbestuiving in een gebied liet zien dat ten minste twee bezoeken nodig zijn voor de maximale zaadzetting in dat gebied. Effectiviteit werd het meest onderzocht door het volgen van insecten. Bij elk kenmerk had een ander insect de meest gunstige waarde. Een fluorescerend poeder (= stuifmeel analoog) experiment liet zien dat stuitmeel in twee dagen 64 meter vervoerd kan worden. Voor A. sylvestris werden dezelfde methoden gebruikt. De hoofdbezoekers (transect waarnemingen) waren Eristalis tenax/pertinax en Empis tessellata. Volgens plot observaties was E. tessellata de hoofdbezoeker. Bij elk kerimerk had een ander insect de meest gunstige waarde. Fluorescerend poeder liet zien dat stuifmeel meer dan 100 meter kon worden vervoerd in een dag. Insecten leken over het algemeen geen speciale voorkeur te hebben voor fluorescerend gekleurde schermen. Op een andere algemene schermbloemige, Aegopodium podagrana, was dehoofdbezoeker Helophilus sp.. Anthriscus sylvestns en hermafrodiete G. hederacea hebben meestal halfgespecialiseerde en vrouwelijke G. hederacea heeft meestal gespecialiseerde bezoekers. De resultaten laten zien dat elke insectensoort haar eigen bijdrage levert aan kruisbestuiving: ze zet veel conspecifiek stuifmeel at, vliegt grote afstanden tussen twee opeenvolgendebezoeken, enz.. Als de relatieve effectiviteit wordt gebruikt en gecombineerd met relatieve voorkomen van de bezoekers, waren Syrphidae rest en Bombus pascuorum de meest belangrijke bestuivers van vrouwelijke G. hederacea. Rhingia campestns was de belangnjkste bestuiver voor hermatrodiete G. hederacea. Erist a/is tenax/pertinax was de belangrijkste bestuiver van A. sylvestris. 5
1. Introduction In Northwest Europe, insects pollinate 80% of the flora. This partof the flora is said to be entomophilous (Kwak, 1 994a). In restoration ecology, it is important topredict possible reestablishment of plants in presence of the available pollinators. Someimportant pollinating insect species could fall out in the case of an environmentalchange (Kwak, 1 994a). Therefore, more information is needed on the relative importanceof insect species for existent plant populations. Mostly, a number of insect species act as the functional groupof pollinators, called the pollinator guild. In some plant communities, keystone species forpollinators may be assigned (Memmott, 1999). A plant species can be visited by a numberof insect species. Memmott (1999) has proposed a prototype foodweb. This foodweb is based oninsect visitation. The total visiting insect pool is divided over the available flowering plant species.If the pollination effectiveness of the different insect species is introduced as a factor into thisvisitation web, a real pollination web can be made. Insects classified as keystonespecies in the food web may play a second fiddle as pollinators. Olsen (1997) defines pollinator effectiveness as the number of seedsthat are 'produced' after a single visit. This depends on the number of conspecificpollen grains that individuals transfer (Herrera, 1987). Effectiveness depends on uptake, deposition,flight distance, foraging speed and flower constancy. Pollinator importance is defined aspollinator effectiveness x relative abundance of visits. Effectiveness of insect guilds of different plantspecies and in different areas can be calculated and compared. This makespossible predictions about seed set. A small population may have to cope with a heterogeneouspollen deposition, because the bulk of the surrounding flowers are of another species,competing for pollinators (Kwak et al., 1998; Heinrich, 1979). Flower constancy of individual insectsplays a role in the rate of heterospecific pollen deposition. The main question in this project is, whether visitation andpollination by insects differ in two morphologically and taxonomically different plantspecies: ground ivy (Hondsdraf), Glechoma hederacea (Lamiaceae) and cow parsley (Fluitenkruid), Anthriscussylvestris (Umbelliferae). The main question is divided into three sub-questions: a.) Which insect species and how many of them are visitors? b.) How effective are these insect visitors separately and as a group aspollinators? c.) What are the consequences of insect visitor pools andtheir effectiveness for pollination? In The Netherlands, the two chosen plant species are often seen atstraight elements in the landscape, like road verges. Road verges are mown several times per year.This mowing regime can influence the number of flowers at a timeand the microhabitat for pollinating insects. G. hederacea flowers from April until June (Van der Meijden,1996). It is a gynodioecious species (Weeda et al., 1988): plants have either hermaphroditic orfemale flowers (Proctor et al., 1996). Both hermaphroditic and female plants can reproduceclonally. The hermaphroditic plants are self-compatible (Widen & Widen, 1990).The oldest flowers at the base are in the female phase (Proctor et al., 1996). Female plants have no anthers andhave smaller flowers than the hermaphroditic plants (Van der Meijden, 1996; Widen & Widen, 1990). Themorphology of Lamiaceae makes them suitable for eutropous (specialised) visitors asbees, bumblebees and butterflies (Ellis & Ellis-Adam, 1994). In the following, Apoidea is used todenote bees and bumblebees. Lepidoptera will be used to denote butterflies. Visitors (andpollinators) on G. hederacea are amongst others bumblebee queens and workers, honeybees, Lepidoptera and Syrphidae (Widen & Widen, 1990; Kwak & Tieleman, 1 994c; Proctor et al.,1996). Bombus spp. (Hymenoptera, Aculeata: Apoidea) live in nests (Proctor etal., 1996). The bumblebee workers start in May (Kwak & Tieleman, 1994c). Queenbumblebees eat pollen and nectar (Kwak & Tieleman, 1994c). Workers need pollen to feed the larvae(Heinrich, 1979; Kwak & Tieleman, 1 994c). The rest of their lives, they need nectar only.Queens emerge in spring, followed by workers. Bumblebees fly during light and avoid rain. A. sylvestris has its flowering season from May to June (Van de Meijden,1996). It is often found together with G. hederacea (Weeda et al., 1987). Umbelliferae are mostlyprotandric and self- pollination is common (Ricciardelli d' Albore, 1986; Weeda et al., 1987). Themorphology of Umbelliferae makes them suitable for allotropous (unspecialised) visitors (Ellis &Ellis-Adam, 6
1994). Visitors (and pollinators) of Umbelliferae are Brachycera, Syrphidae, Apoidea, wasps, Formicoidea, Coleoptera, and Hemiptera species, which are mostly allotropous insects (Koul et al., 1993; Grace & Nelson, 1981; Ellis & Ellis-Adam, 1994). Syrphidae and Coleoptera are hemitropous (half specialised) insects (Ellis & Ellis-Adam, 1994). Scatophagidae can be expected as visitor of A. sylvestris (pers. Hotfmann). Scatophagidae arepredators, especially in wet biotopes (Van Veen, 1993). 7
2. Materials and methods Appendix A gives a research calendar. In this calendar, exact dates and areas of the investigations are given. 2.1 Background information Seed set Crossing experiments were done in order to obtain information about the overall effectiveness of pollination by the available insect guild. From the handcross-pollination, maximum seed set per fruit can be calculated (Herrera, 1987). Plants werenetted and kept in an experimental garden to let them develop seeds under similar conditions. From those data canbe concluded if insects are necessary for pollination. To investigate the breeding system, five hermaphroditic and five female G.hederacea clones were in triple either not (or hand self) or hand cross-pollinated at 9 a.m., 9 a.m. +3 p.m. or 3 p.m. in the greenhouse. We used potted plants that had been keptin the experimental garden. Hand-pollination in the field is a kind of control on resource availability (Widen & Widen, 1990) when compared to natural insect pollination. Seed set of G. hederaceaflowers that had been open (insect) pollinated in the field were compared to seed set of flowersthat had been extra hand cross-pollinated in the field. Hand pollination was performed byputting an anther against a stigma (Corbet, 1999). Twenty days later, after full ripening of the seeds,seeds were collected. Seven potted A. sylvestris plants were hand pollinated by rubbing a male umbellule against a female umbellule. Whole umbellules were either not or hand self orhand cross- pollinated at 9 a.m., 9 a.m. + 3 p.m. or 3 p.m. in the greenhouse. Every plant had seventreated umbellules. The plants were kept in the experimental garden. More than two weekslater, after full ripening of the seeds, seeds were collected. Nectar production, attractiveness and pollen presentation Nectar contents of flowers of eight netted G. hederacea plants (four hermaphroditic and four female) were measured at 10 a.m., noon, 2 p.m. and/or 4 p.m. with amicrocapillary of 1 microliter. Production by G. hederacea was compared with Lamium album, a speciesthat may be competing or facilitating. Nectar contents of flowers of eight netted L. album plants were measured at 10 a.m., noon, 2 p.m. and/or 4 p.m. with a microcapillary of 2 microliter.Nectar production at different times of the day was calculated. With a field refractometer,the sugar concentration in the nectar was measured. These data show at what time of the day the nectar standing crop is favourable for visitors.
2.2 Visitors Verge composition Plant composition in the verges was recorded by counting the number of flowers(dates and verges: see appendix A). The data can give insight in patterns of flowerdiversity and the possibility for flower choice for insects.
Plots Three areas with a difference in plant and insect species richness were chosen as research areas (Appendix B). The research was carried out in three areas nearAssen in the province of Drenthe in the northern part of the Netherlands (52°56' N - 53°45' N,6°27' E - 6°41' e) (figure 1). Stroomdal Drentse Aa is a brook valley. Most of the area is inagricultural use (Prins et al., 1998). Bovensmilde is the most intensive agricultural area, whereasDrentse Aa is the least intensive agricultural area. Drentse Aa is located next to the nature reserve 'Drentse Aa'.The third area, with intermediate agricultural use, was Ekehaar. In plots, numbers of visits of all insect species were recorded during ten minutes.Plot observations were made at different times of the day. Recordings contain: insect species, residence time, number of visited inflorescences or umbels, and number of visited flowers or 8 umbellules. Also the number of available infiorescences orumbels and total available number of flowers or umbellules in the plot were recorded. Fromthose recordings, number of visits per insect species, visited inflorescences or umbels andduration of visits were calculated. With these data, information about insect guild composition (duringthe day), foraging speed, and insect behaviour concerning effectiveness (see results) wereobtained.
A. The flerIands.v' sohm B.Lrenhe
— road povind.I kwdatj enlarged river vilkye orOW1) z vesyenwnber
Figure 1. Research area. A. The Netherlands. B.Drenthe, a province In the north-east of The Netherlands. C. The research area in the surroundings of Assen with Itsposition of verges. Verges 25, 251/2, 26, 26'/z, 27, 27½, 34, and 35 belong to Drentse Aa. Verges 17, 17½, 18, 18½, and 191/2 belong to Ekehaar, which is an area with slightly more Intensive agriculture. The most intensive agricultural area is Bovensmilde with verges 2, 5, 9, 12 and 40. Road verge numbers refer to the ones used by F. Hoffmann. 9
Transects By transect observations, the overall compositionof the visitor (and pollinator) guild can be estimated (Dramstad, 1996). Visiting insectspecies were spotted while slowly walking the transects. The duration of the walks, the numberof insects per species and the available number of inflorescenceS or umbels were recorded. Thelength of the transect was equal to the size of the population. 2.3 Effectiveness of visitors Following insects The choices of the bumblebee species as a whole aredefined by the choices of individual bumblebees, especially bumblebee workers canhave temporal flower constancy (Kwak & Tieleman, 1 994.c). A comparable situationcould be the case in A. sylvestris. With help of a stopwatch, total visiting time was recorded. The flight distancesbetween two inflorescences or umbels were estimated. The foraging speed wascalculated. The number of visited flowers per inflorescence or number of visited umbellules per umbel(a measure for efficiency), and total number of flowers or umbellules were recorded of visitors onboth plant species. Their average forage speed, foraging efficiency and flight distance werecalculated.
Pollen loads Insect visitors can take up pollen from the anthersduring a visit. The total number of pollen grains of different plant species an insect bears,is called the pollen load. Visitors were slightly anaesthetised by applying CO2 with a Corkmaster pen.The pollen loads were removed with a sticky staining gel (Beattie 1972). The colouringagent of the gel was basic fuchsin. The gel was melted in order to makepreservable slides. Slides were analysed microscopically (10 x 40). Pollen grains were identified morphologically (shapeand size) and counted. Percentages of target pollen, number of pollen species and totalload were derived. From these data, size of pollen loads of insect species and their flower constancy wereobtained. Visiting the flowers of Lamiaceae, pollen is deposited on thedorsal side of bees (Weeda et al., 1988). Pollen were seen between the eyesand on mouthparts. For that reason, pollen was only removed from the back and the heads of visitorsof G. hederacea. If present, corbicular loads were not collected, because those pollencould not longer pollinate flowers. Pollen was removed from the ventral side of visitors of A. sylvestris.
Pollen deposition Flowers of netted (unvisited virgin) G. hederacea plants wereemasculated. Visitors were allowed to visit in a sequence three virgin pistils at most.The stigmas were melted into sticky staining gel. Numbers of pollen per plant species in thepreparations were counted as described in 'pollen loads'. Total deposition and percentageof target pollen were calculated. From the data information about flower constancy was obtained.
Fluorescent dye powder The distance of pollen transfer by the pollinatorguild was estimated by applying fluorescent dye powder (Radiant Colour NV) on a source groupof plants and by collecting plant parts at different distances from the source(Kwak, 1 994b). The stigmas were melted into a colourless gel (Beattie, 1972). The fluorescent dye particlesin the preparations, grouped as red, yellow and green, were counted under a microscopewith UV-light (10 x 40) in a dark room. From these data, the overall pollen transfer distance wasobtained. For G. hederacea, fluorescent colours CLEAR (R09; batchnumber M3252) and CERISE (RC1 6; batch number 3514) were put onto 10 hermaphroditicflowers. The next day, ten flowers were collected at distances 0, 1, 2, 4, 8,16, 32 and 64 meters west of the source and 10 and 16 meters east of the source. Fluorescent colour CLEARconsisted of yellow and green particles as seen under the UV microscope. Fluorescentcolour CERISE consisted of red particles as seen under the UV microscope. For A. sylvestris, fluorescent colours CLEAR, CHARTREUSE(RC1O; batch number 3595, ref. 6348), ORANGE (RC13; batch number 3262, ref. 6348),and CERISE were put on eight x two umbellules on the same umbel, each colour on a differentumbel. At the end of the 10 day, ten umbellules per distance were collected at distances 1, 2, 3 and 4 meters north and at 4, 5, 10, 20, 40, 75 and 100 meters south of the source. Twenty umbellules were collected near the source. Fluorescent colour CLEAR consisted of green and yellow particles as seen under the UV microscope. Fluorescent colour CHARTREUSE consisted of green particles as seen under the UV microscope. Fluorescent colour ORANGE consisted of yellow particles as seen under the UV microscope. Fluorescent colour CERISE consisted of red particles as seen under the UV microscope. To contro' preferences of A. sylvestris visitors for fluorescent dye colours, visitation on umbels of another umbellifer flowering abundant at that moment, Aegopodiumpodagraria,was observed. The colours CERISE, ORANGE and CHARTREUSE were put on one side of the umbels of seven umbels. The other sides were used as controls, and there were seven blank umbels. The data give information about the visitor guild of A. podagraria(populationAssen) and of the preferences of the visitors for the differently coloured umbels. Statistics SPSS was used to analyse the data statistically, mainly by ANOVA and Tukey tests. 11
3. Results The results of G. hederacea and of A. sylvestris can be foundin chapter 3.1 and 3.2 respectively. 3.1 Glechoma hederacea- Background information In this chapter, the breeding system of G. hederacea will bepresented, together with extra hand pollination in the field measurements. Raw data canbe found in Appendix E (G. hederacea breeding system experimental), Appendix F (breeding systemG. hederacea field), Appendix H (nectar G. hederacea) and Appendix I (nectar L.album).
Verge composition Differences in visiting species, number of visits and visitor rate canbe due to the environmental conditions. The abundance of flowers of different plantspecies in a verge can be important for the attractiveness of a verge. Also can different vergecompositions provide different habitats and food sources for visiting insects. In Bovensmilde, 9 flowering plant species were observed,whereas in Ekehaar and Drentse Aa respectively 5 and 6 flowering plant species wereobserved. The average number of flowering plant species in Bovensmilde was 3.57; The averagenumber of flowering plant species per verge in Ekehaar was 2.33. The averagenumber of flowering plant species per verge in Drentse Aa was 2. The average number of flowers in Bovensmilde(per verge) is 733; The average number of flowers in Ekehaar was 2015;the average number of flowers in Drentse Aa is 102. Ekehaar had the most flowers per verge, whereas DrentseAa had the least flowers per verge. The average number of G. hederacea flowers per verge was452 in Bovensmilde, in Ekehaar 1507 and in Drentse Aa 68. There were large differences in number of flowering plant speciesand number of flowers in the different areas. Drentse Aa would be expected to bevisited most, because in 2000, it had the largest number of flowering plant species (Hoffmann,unpublished data). Other attractive plant species for bumblebees were L. album and Lamium purpurea,Silene dioica and Symphytum officinale. Those plant species could compete with or facilitate forvisitors on G. hederacea.
Breeding system Glechoma hederacea To make a good estimation about the influence of behaviour andflight distance, information about the breeding system is needed. Experimentallyobtained data are given in figure 2. Female flowers even produced seeds without pollination: 0.4seeds per flower. Somehow pollen must have reached the stigmas. They produced moreseeds after cross pollination. Cross pollination at 3 p.m. had more effect than at 9 a.m. (1.7 and 1.2respectively). Cross pollination at 9 a.m. and 3 p.m. resulted in the highest number of seeds(1.9). The hermaphroditic type did not produce seeds withoutpollination. With self pollination at 9 a.m. and at 3 p.m., 1.3 and 1.0 seeds are producedrespectively. When flowers were double self pollinated, the seed set was even higher (1.5). Crosspollination at 9 a.m. and 3 p.m. resulted in equal amount of seeds (1.2). double cross pollination at 9 a.m.and 3 p.m. resulted in a higher seed set of 1.9 seeds per flower. Overall, cross pollinationresulted in a higher seed set than self pollination. The female type has a higher seed set than thehermaphroditic type, but this difference is not significant. Double cross pollination resulted insignificantly higher seed set than no pollination in hermaphroditic plants. 12
Table 1. Flowers in verges. Of all flowering, insect visited plants in the investigated verges, the number of flowers was counted. Column 1ives the date in the order: year-month-da" '3
2. Breeding system ofGlechoma hederacea 0femaleu hermaphroditic
14 14 1512 15 14 n= 1510 -12 -8 -13
2.5
2 0 1.5
1
0.5
0
Pollination treatment
Figure 2. Breeding system of Glechoma hederacea. in the greenhouse, bagged flowers wereself or cross single pollinated at 9 a.m. or 3 p.m., or double pollinated at 9 a.m. + 3 p.m.Self means self pollination, cross means cross pollination. The numbers on the x.axis are times inhours. 9 means 9 a.m. 15 means 3 p.m. 9 + 15 means 9 a.m. and 3 p.m. Y error bars arestandard errors from the means. The average number of seeds per flower isgiven. N is the number of flowers per treatment. Homogeneity of variances Is 0.000. The chance forequality of means (ANOVA) Is 0.019. the Tukey test divides the means in two groups: a.with a chance of 0.078 and b. with a chance of 0.194. 14
Field extra pollination Extra pollination in the field can reveal if a low seed set is due to alow visitation or pollination rate. When the pollination activities are not the bottleneck,equalseed set between treated and control is expected.
3. Seed set of Glechoma hederacea in thefield
Dextra hand pollination U control
C) 0 I
hermaphroditic Plant type
Figure 3. Seed set of Glechoma hederacea with extra handpollination. In verge 34, female and hermaphroditic flowers were extrahand pollinated. The controls were flowers In the same flowering stage on the sameplant. N is the number of investigated flowers. Mean plus standard error Is shown. A paired-sample ttest concluded that the controls did not differ from the treatments (signIficance 0.812).There Is also an equality of means of the controls of different plant types, with asignificance of 0.394. The means of the treatments of the different plant types areunequal, with a significance of 0.017.
Hermaphroditic plants had both in the natural and in the treated situationsignificantly more seed set than female plants (2.6 and 2.4 and 1.3 and1.8 respectively). Hermaphroditic plants had a higher seed set after extra hand pollination (2.7 instead of 2.3, notstatistically significant), whereas female plants had a lower seed set after extra handpollination (1.3 instead of 1.8, not statistically significant). 15
Nectar production rate by Glechomahederacea Differences in visitation of female andhermaphroditic G. hederacea could be explained by differences in nectar production of the twoplant types. Plants were kept in pots in a greenhouse during this experiment. Female and hermaphroditic flowers did not differin nectar production rate by an ANOVA (Appendix C) so data were pooled. Figure 4 shows,how the nectar production speed diflers during the day and under different treatments.Appendix H shows a concentration of 30 to 50%.
4. Nectar production by Glechoma hederacea
0.06
L. 47 63 29 33 n=26 6a2 2: a a 5 0.05
C., E 0.04 0 I- w a. 0.03 C 0 4- C., 0.02 0 a.
II, IrTh 0 2 9 2 9 2
Treatment
Figure 4. Nectar production per hour byGlechoma hederacea. Treatment 10 means 10 a.m., 12 means noon,14 means 2 p.m., 16 means 4 p.m. Nectar was depleted at 4 p.m. theday before the nectar amount measurements. There areten 'treatments', In which the production periods andproduction times varied. Means plus standard errors are shown. N is the number ofinvestigated flowers. The homogeneity of variances is 0.000. The chance that the means comefrom the same population Is 0.002 (ANOVA). The means can be classified into two groups(Tukey). The probability of group a. Is 0.686. The probability of group b. is0.110.
The average production speed per hour was highestin treatment '12 to 14': 0.027 microliter per hour, but the sample size was only 2.Other treatments during the day resulted in nectar production from 0.012 to 0.0083 microliter perhour. Overnight treatments showed a slow night production rate: values were between 0.001 and 0.003microliter per hour. Overnight values differ statistically significant from most of the valuesduring the daytime. 16
Nectar production rate by Lamium album In some verges, L. album was flowering. Nectarproductionrate of L. album can be compared with production of G. hederacea.Thedifferences could elucidate attractivity differences of verges with or without L. album.
5. Nectar production by Lamium album
0.6
0.5
0.4 0 .c 0. 0.3 C 0 (3 0 0. 0.1 C, z 0
Treatment
Figure 5. Nectar production rate by Lamium album The nectar production is calculated from nectar contentsIn plants kept in the greenhouse. Nectar was depleted at 4 p.m. the day before the measurements.Treatment 10 means 10 a.m., 12 means noon, 14 means 2 p.m.,and 16 means 4 p.m. There are ten treatments with different production periods. Treatment 10 to 12 means that it wasemptied at 10 am. The nectar content at noon gives the production in the two hours.Of each period, the nectar production per hour was calculated. Means plus standard errors areshown. N Is the number of flowers. The homogeneity of variances is 0.000. Thechance for equality of means is 0.000. Tukey divides the means into two groups: a.with a chance of 0.172 and b. with a chance of 0.949.
Per hour, most nectar was produced at the treatment '12 to 14': 0.39microliter. The other treatments at the daytime resulted in production per hour of 0.25 to 0.32microliter. Overnight treatments resulted in statistically significant lower production of 0.03 to 0.06microliter per hour. 17
Sugar concentration of Lamium albumnectar For visiting insects, not only the amount of nectar,but also the sugar concentration in the nectar is important. Figure 6 shows sugarconcentration of L. album nectar.
6. Sugar concentration of Lamium album nectar
n= 7 7 10 4 7 5 17 10 11 1 abc abc ab bc a bc bc C C 50
45 Q
0
25
20
15
10 (I) 5
0 2 .9 9 2 9 2 9 2 2 2 .9 0 0 0 C1 CD CD CD CD
Treatment
Figure 6. Sugar concentration of Lamium albumnectar. With a refractometer, the concentration wasmeasured in nectar produced at the different treatments of figure 5. N Is the number ofinvestigated flowers. Means plus error bars are shown. The homogeneity of variances is 0.103.The chance for equality of means(ANOVA) is 0.000. Tukey divides the means into group a.with a chance of 0.080, group b. with a chance of 0.190 and c. with a chance of 0.776.
The highest sugar concentrations (42.5 to 46.2%) werefound in the 'overnight' treatments. Sugar concentrations of 28.2% to 42.3% werefound in the treatments at the daytime. The lowest sugar concentrations were found inthe nectar of '10 to 16' (32.1%) and '12 to16' (28.2%). '14 to 16' on the other hand, had a concentrationof 41.6%. 18
3.1Glechomahederacea- visitors Plots Todetermine the composition of the visitor guild, plotobservations were done (see 2.1.1). The results are presented in figure 7. Raw data canbe found in Appendix J (plot G. hederacea). Both female and hermaphroditic plots wereobserved. Figure 7 shows the average of the fraction of visited flowers per plot by differentvisiting insect species. The total number of flowers was estimated by counting the number offlowers on 25 randomly chosen inflorescences. This resulted in an estimation of the average numberof flowers per inflorescence. The total number of flowers in a plot was calculated bymultiplying the total number of inflorescences per plot and the average number of flowers per inflorescence.
7. Fraction of visited Glechoma hederacea flowers
0 female •hermaphroditic
b b b b ab a b ab
n=8 8 9 6 8 4 3 1 0.020 w .0 0.016
0 I-(n WQ)o0.012 = 0.008
0.004
0.000 Melanostoma Rhingia campestris Apis mellifera Bombuspascuolum spiPlatycheirus sp. Insect species
Figure 7. Fraction visited Glechoma hederacea flowersof total available flowers in a plot. Every plot observation took 10 minutes. 18 doubleplot observations were done in verges in the three areas. Per plot, the fraction visited flowersof the total number of flowers was calculated. N is the number of observed insects. Mean valuesplus standard error Is shown. A one-way ANOVA was done to compare the means.After a log-transformation, the homogeneity of variances was 0.000. The significanceof the ANOVA was 0.005. The values can be divided in two groups: a and b (Tukey test).Tests were done per insect species female and hermaphroditic. 19
The fraction of visited flowers was for all insect species higher on hermaphroditic plots than on female plots. Apis mellifera took a larger fraction in both plant types than the other insects: 0.009 on female plots and 0.014 on hermaphroditic plots. Bombus pascuorum had also a high fraction of visited hermaphroditic flowers (0.008), but this value does not differ significantly from visitation fraction values of the other insects (ANOVA). Most of the visits on both female and hermaphroditic flowers were made by A. mellifera, which was thereby an important (frequent) visitor. On hermaphroditic plants, B. pascuorum was also an important visitor, but this is not significantly more important than visits by Melanostoma sp.IPlatycheirus sp. and Rhingia campestris. Hermaphroditic plants attracted more visits than female plants, though this is not a significant difference. The overall fraction of visitation on female flowers was 0.0118. The overall fraction of visitation on hermaphroditic flowers was 0.0293.
Transects Besides plot observations, transect observations were done. Comparisons of results of those two methods can elucidate the differences between the methods. In this way, wemight even compare plot observations of one species with transectobservations of another species, after minor transformations. This could be used in comparative studies in which plant specieswith different visitor characteristics are used. Transect observations on female and hermaphroditic verges were kept apart. Every walking transect observation, the number of inflorescences was estimated. In total, 4839 female plants and 13566 hermaphroditic plants have been observed. Figure 8 shows the results. Raw data can be found in Appendix L (transect G. hederacea). 20
8. Visitors on Glechoma hederacea
D Lepidoptera n=15 n=47 50 • Enstalis sp. IIIIIlIllIIIIIIIIII o Helopliilus sp. 40 III • Meianostoma spiPlatycheirus sp. 3o o Rhingia campestns
C •Diptera small .E20 (0 • Apis mellifera > IlllilüiUhlI. 10 o Bombus terrestris
o Bombus pascuoium 0 female hermaphroditic Planttype
FIgure8.Visitors on Glechoma hederacea, derived from transect observations. Transect observations were made in all areas. The number ofobserved insects In total during the transect walks is given per plant type. On female G.hederacea, 15 insects were observed, whereas on hermaphroditic G. hederacea, 47 Insects wereobserved. In total, transect observations were made along 4839 female and13566 hermaphrodItic plants. Female plants have on average 3.72 flowers per inflorescence.Hermaphroditic plants have on average 2.68 flowers per Inflorescence.
At the female G. hederacea, in total 13/4839 = 0.0027 visitors perinflorescence were observed, whereas at the hermaphroditic in total 47/13566 = 0.0035visitors per inflorescence were observed. Female and hermaphroditic plants had almostthe same number of visiting insects. The composition of the visitor guild differed: main visitors atfemale G. hederacea were A. mel/hera and B. pascuorum,whereasthe main visitor at hermaphroditic G. hederacea was R. campestns. A. me/lifera and the butterflies were not observed onhermaphroditic G. hederacea at all. R. campestris did not visit female G. hederacea. Female andhermaphroditic plants had relatively the same amount of visitors. (4839/13 is approximately 13566/47). 21
Area Transect observations were done in the three different research areas.The total number of visits made by visitors is compared in the three areas.
9. Visitors on Glechoma hederacea inthree differentareas
n transect observations = 2 10 1 0.40
C) 0. 0.35 C) U C C) V.) 0.30 'a, C) L. 0 0.25 C a 0.20
V 0.15 > V C 0.10 ci C
'4, 0.05 > 0.00 Bovensmilde Ekehaar Drentse Aa Area
Figure 9. Visitor pressure on Glechoma hederacea in threedifferent areas. The number of average visiting individuals per verge,obtained from transect walks is given. N is the number of followed Individuals. Meansplus standard error Is shown. The homogeneity of variances is 0.000. According to the one-wayANOVA, the significance for equal means is 0.256. The chance that the means belong tothe same group is 0.439 (Tukey). The differences are not significant at the a = 0.05level.
In Bovensmilde, the average number of visiting individuals perinflorescence per verge was 0.02, in Ekehaar it was 0.01. Inthe Drentse Aa, the chance for finding an insect on an inflorescence was 0.34. The differences are not significant. 22
3.1 Glechoma hederacea — effectivenessof visitors Some traits that make one insect species more effective as apollinator than another are pollen loads, pollen deposition, efficiency of behaviour,foraging distance arid flower constancy. Data of those traits will be presented in this chapter. Rawdata are presented in Appendix N (following G. hederacea) and Appendix J (plot G. hederacea).
Flight distance The further away the next visited inflorescence, the higherthe chance of cross pollination. Figure 10 shows flight distances derived from data fromplot observations.
10. Flight distances of Glechoma hederacea visitors
30
w 0 () 0 25 C, Ce 0 20 C C h15
0 C, C 10 4-Ce 0Ce 4- 5
U.
0 Melanostoma Rhingia campestns Apis mellifera Bombus sp sp.JPlatycheirus sp. Insect species
Figure 10. Flight distance of Glechoma hederacea visitors. The data were obtained by plot observations. Distances wereestimated. N Is the number of observed insects. Means plus standard errors are shown. Flightdistance is defined as flown distance in cm between two visited Inflorescences. 21 insectsspread over four Insect species were observed. The homogeneity of variation is 0.143.The significance for equality of means is 0.631 (ANOVA). According to Tukey, the significancefor equality of means is 0.795. 23
R. campestris had the highest average flight distanceof 18.5 cm, followed by Bombus sp., 11.2 cm. The lowest averageflight distances were observed for Melanostoma sp.IPlatycheirus sp. (11.0 cm) and A. mellifera (10.2 cm). Table 2 shows the maximum flight distances of thevisitors. This could be an important measure for potential efficiency.
Table 2. Maximum flight distance between Inflorescencesof Glechoma hederacea visitors
Insect species Melanostoma sp. Rhingia Apis mellifera Bombus sp. /Platycheirus sp. campestns
Maximum flight distance (cm) 18 45 13 35
Visitation speed As described in 'transect observations', transectobservations could be converted into plot estimations by using the visitation speed of thevisitors. Figure 11 shows the visitation speed of the main G. hederacea visitors. The data arecollected by following individual flower visiting insects. Because the data of visitors on female G.hederacea did not differ significantly from the data of visitors on hermaphroditic G. hederacea flowers, nodivision was made in the visited plant type.
Bombus sp. had the highest visitation speed of 18.9 flowers perminute, followed by A. mellifera, which visited 13.6 flowers per minute. Both hoverflies had significantly lower visitation speeds: Melanostoma spJP!atycheirus sp. visited 3.5 flowers perminute, and R. campestris visited only 3.1 flowers per minute. 24
11. Visitation speed of Glechomahederacea visitors data from following observations
25 o 20
C E 015 0. I-U) 0 010 •0 0 >5(a)
0 Bomous pascuorum MelarioStOma Rhingia campestris Apis mellifera sp./PiatycheiruSsp. Insect species
Figure 11. Visitation speed of Glechoma hederaceavisitors. The number of visited flowers per minute wasderived by folowlng individual flower visiting individuals. N is the number of observedInsects. Means plus standard errors are shown. Visitation speed is defined as timefollowed divided by the total amount of visited flowers. We followed In total 25 indivIduals spread overfour Insect species. The homogeneity of variances was 0.017. An ANOVAshowed a significance for equality of means of 0.000. The Tukey test showed adifference between A. mellifera and Bombus sp. on one hand (signifIcance 0.999)and Melanostoma spiPlatychelrus sp.and R. campestris on the other hand (significance0.448). This graph takes all protocols intoaccount with three or more flowers visited, because if only one ortwo flowers are visited, the visitation speed can not be calculated. 25
Visitation speed from plot observations The data of figure 11 were derived from following individual flowervisiting insects. Actually, during plot observations, visitors were also followed. Comparisonof the figures can be interesting and elucidate mistakes during the research: following was onlydone in a few verges.
12. Visitation speed of Glechoma hederacea visitors, data from plot observations
E
Melanostoma Rhingia campestris Apismellifera Bombus pascuowm spiPlatycheirus sp. Insect species
Figure 12. Visitation speed of Glechoma hederacea visitors. Data are derived from plot observations. N Is the number ofobserved insects. Means plus standard errors are shown. The homogeneity of variances is 0.555.The chance for equality of means is 0.806 (ANOVA).
The number of visited inflorescences per minute is taken as a measure forvisitation speed. Because in each inflorescence at least one flower was visited, thevalues were expected to be higher than in figure 11. Bombus pascuorum had the highestvisitation speed: 11.8 inflorescences per minute. Apis mellifera and R. campestris were intermediate.Melanostoma spiPlatycheirus sp. had the lowest visitation speed of 0.089 inflorescences per minute. Though differences are not significant, B. pascuorum seems to be the fastest visitor, whereas Melanostoma sp./P!atycheirus sp. was the slowest visitor. All insectspecies, except for R. campestris, had a higher value for the number of flowers per minutethan for the number of inflorescences per minute (see figure 11). Values of inflorescences per minute can be transformed by dividing the number of visited flowers per minute by the average number of flowers per inflorescence an insect species visits. Those values are given infigurel 3. 26
Effective behaviour Another characteristic of the visitors is their tendency to visit many or few flowers per intlorescence. Data were obtained both by following individual flower visiting insects(figure 13) and by plot observations (figure 14).
13. Percentage visited flowers per Glechomahederacea inflorescence data derived from following observations
40
L. 35 0. U) 30 C, OC) 25
20 >0 15 .5.- C, U 10 C) 0. 5
0 Melanostoma Rhingia campestris Apis mellifera Bombuspascuorum spiPlatycheirus sp. Insect species
Figure 13. Percentage visited flowers of presented flowers on the visitedinflorescence by Glechoma hederacea visitors. Data are derived from following individual flower visiting Insects. N is the numberof observed insects. Means plus standard errors are shown. Presented flowers isdefined as total number of flowers on visited inflorescences. 15 insects of four species were followed. The homogeneity of variances is 0.379. Differences are not significant(ANOVA, p= 0.380)
Rhingia campestris visited the most flowers from a presented inflorescence (= 25.4%). Apis mellifera, B. pascuorum and Melanostoma sp./Platycheirus sp. had percentages of 10.2 to 11.6. The differences are statistically not significant (ANOVA). 27
14. Percentage visited flowers perGlechoma hederacea inflorescence data fromplotobservations
0 female U hermaphroditic
ii' mellif era Bombus pascuorum Melanostoma Rhingia campestris Apis sp./Platycheirus sp. Insect species
hederacea Figure 14. Percentage visited flowersof presented flowers by Glechoma visitors. N is the number of observedinsects. Means plus Data are derived from plot observations. number of flowers per standard errors are shown. Presentedflowers is defined as the total visited Inflorescence. Of four specIes, 47Insects were followed. Thehomogeneity of variances was 0.266. An ANOVA showed asignificance of 0.099 for equality of means. visited flowers. The There are no differences betweenthe Insects in their percentages overall mean is 44%. visited. A percentage of 100 means, that allflowers of the visited inflorescenCeS were and of 55 on Bombus pascuorum has the highest percentage,of 58 on female inflorescenceS hermaphroditic inflorescences. MelanostomaspiPlatycheirLlS sp. had the lowest percentage hermaphroditic (37.5) on female flowers, whereas R.campestris had the lowest percentage on at the female inflorescenceS (28). For all insect species,the percentage was somewhat higher inflorescenceS than at hermaphroditic inflorescenCeS. 28
Pollen loads The carrying capacity of a visitor is given by itspollen load. Not only the number of grains, but also the purity of the load are importantfor the pollination of the receiving flower.Raw data can be found in Appendix P (pollenload preparations G. hederacea).
15. Pollen loads of Glechomahederacea visitors 0 Glechoma hederacea pollen • rest pollen
5000
4000
2000
1000
0 Bombus sp. Melanostoma Rhingia campestris Apismellifera spiP!atycheifuS sp. Insect species
Figure 15. Pollen loads on Glechoma hederaceavisitors. The pollen in the brushed pollen loads of visitinginsects were classified Into G. hederacea pollen and other pollen. N Is the number ofbrushed Insects. Means and standard errors are shown. Comparison of means wasdone by ANOVA tests. Tukey tests weredone to group the means. The variance of thenumber of G. hederacea pollen could notbe made homogeneous. The significance of the ANOVA wasfor the 6. hederacea pollen 0.070 for the total number of pollen 0.000 and for the numberof pollen species 0.429. The Tukey test was done to group the means of thetotal number of pollen. Three groups canbe made: a. R. cam pestris and A. melllfera (0.996); b. Apis melliferaand Melanostoma spiPlatychelrus sp.(0.073) and c. Melanostoma spiPlatychelrus sp. andBombus sp. (0.502)
Bombus sp. had the largest load: 1623 on the female typeand 4568 on the hermaphroditic type. Rhingia campestrishadthe smallest pollen load of 62, followed byA. mellifera on female (120) and hermaphroditic (159) G. hederacea. 29
Table 3 shows other characteristics of the pollen loads of G. hederacea visitors.
Table 3. Characteristics pollen loads of Glechoma hederacea visitors.
Insect species Number of Fraction Pollen load size pollen species Glechoma hederaceagrains
Melanostoma sp./Platycheirus sp. 6.0 0.94 690 Rhingia campestris 5.0 0.69 62 Apis mellifera 6.3 0.47 124 Bombus sp. 6.4 0.87 3387
All visitors had between 5 and 7 pollen species in their loads. The fraction of G. hederacea pollen was highest for Melanostoma spiPlatycheirus sp. (0.94). Thefraction of G. hederacea pollen was lowest for A. mellifera (0.47). The differences betweenBombus sp. and both A. mellifera and R. campestris are statistically significant (ANOVA). Bombus sp. is a visitor with a large load on hermaphroditic (and female) plants. A. mellifera and R. campestris have lower capacities. The load of A. melliferadid also consist for half of other pollen species. A fully loaded Bombus sp. can deposit pollen on more female flowers than A. mellifera (supposed that their number of deposited pollen grains in a visit is the same). Deposition characteristics are shown in figure 16. 30
Pollen deposition Without a load of G. hederacea pollen no pollination is possible. Figure 16 shows the deposited G. hederacea pollen on the stigma of the first visited flower. Because data on second and third pistil could be inaccurate, figure 8 shows only deposition onto the first pistil. Raw data can be found in Appendix A (virgin stigma slides). Appendix 0 gives pollendeposition on the second and third stigma also.
16. Pollen deposition on virgin Glechoma hederacea pistils
n=2 3 12 2 1 13 1 3 3 3
100 90 0 a80 70 U 60 50 40 30 20 10 I 0
Lepidoptera Syrphidae Apidae insect species
FIgure 16. Glechoma hederacea pollen deposition. Data are derived from first visitation on emasculated G. hederacea flowers. N Is the number of first visited pistils. Means plus standard errors are shown. Of slides of visited pistils, the average number of G. hederacea pollen is given.
It is expected, that pistil 1 receives more pollen than the next pistils. This was not the case. See Appendix D. Higher values can be caused by improper emasculation or ahigh variation in deposition. Syrphidae deposited 14.3 grains on the first pistil. Lepidoptera deposited 11.4 grains on the first pistil. R. campestris and B. pascuorum were intermediate with 9.1 and 8.1 pollen respectively. A. mellifera deposited 6 grains on the first pistil. Apidae large and small deposited 3.2 and 4.0 grains respectively. Bombus pratorum and Bombus terrestris (which visits through a self-bitten hole) deposited the low numbers of 2 and 0.7 pollen grains on the first pistil. 31
Flower constancy The more flower constant an insect species is, the less different pollen grain species will be deposited. Figure 17 shows the number of pollen species in the deposits.
17. Pollen species on Glechoma hederacea pistils
• Pistil 1 • Pistil 2 0 Pistil 3
n=3 3 24 2 3 35 3 8 4 7 4.5
4
3.5
U &2.5 0 C 2 o 1.5 0.
0.5
0
Insect species
Figure 17. Number of pollen species on Glechoma hederacea pistils. Data are derived from visitationonstigmas of emasculated G. hederacea flowers. N is the number ofslides.Meansplus standard errors are shown. Of 92 microscope slides of visited pistils, average number of pollen grain species is given.
B.pratorum left most pollen species (4;3;3). The least pollen species were deposited by Lepidoptera (1.3). With A. mellifera, Apidae small, Apidae large, B. terrestris, and R. campestris, the first pistil has received less pollen species than the second or third. The other pollen species could come from flowering species like Lamium album in the neighbourhood. Identification was not possible, but grains with shapes typical of Umbelliferae were not much found. ______
32
Fluorescent dye powder The overall result of the visitation of the whole pollinator guild can be shown by fluorescent dye powder results. They are presented in figures 1 8a, b and c. Raw data are presented in Appendix S (G. hederacea fluorescent dye powder). Green was not much present at the source, but was found in samples at 64 metres (15). To the east was a clear decay of the total number of fluorescent dye particles (17 to 9). To the west, the number of particles seems to be uncorrelated with distance. At 32 meters, three times as many particles were found than at 2 meters (29 and 10respectively). A decay for the colours separately could not be found either.
18a. Red fluorescent dye powder on Glechoma hederacea
—,--.
0.8
0
I, I .2 a V V . —------——-—0.2---- I I f I •.f f II. -----—----—----Oi—- —-'-—---—'— -10 0 10 20 30 40 50 60
0.2 Distancefrom source(meters) 33
1 8b. Yellow fluorescent dye powder on Glechoma hederacea
0 0 0 U U 0 U
tU a. >. 0 C 0 U U I,.
Distance from source (meters)
18c. Green fluorescent dye powder on Glechoma hederacea
U U 0 0 U 0 0 U t0 a.
0 C 0 U U I,.
20 -10 0 10 20 30 40 50 60 70 Distanc. from .ourc. (m.t.rs)
Figure 18 a, b, c. Red, yellow and green fluorescent dye particles on Glechoma hederacea. Data are derived from a fluorescent dye powder experiment. The colours are the colours seen under the UV microscope. Means plus standard errors are shown. The sample location is given in meters. 0.5 meters Is the source. + values are at the west, - values are at the east. 34
3.2. Anthriscus sylvestris- backgroundinformation To make good conclusions about effectiveness of visitors, informationabout the breeding system is needed (figure 19). The overall effect of the visitors isshown in figure 27, about fluorescent dye powder. The control on colour preferences is shownin figure 28. Raw data can be found in Appendix G (breeding system A. syl vest risexperimental).
Breeding system The breeding system has to be elucidated, to make conclusions aboutharmfulness of short flight distances.
19. Breeding system of Anthriscus sylvestris
a. a) In V
In a) 0) C 0a) 0a)
Pollination treatment
FIgure 19. Breeding system of Anthriscus sylvestris. In the greenhouse, netted plants were hand pollinated in different ways.Self means self pollination, cross means cross pollInation. 9 means 9 a.m., 15 means 3 p.m. N Is the number of replicas, this is 7 for each treatment. Means plus standard errors areshown. Percentage seed set is defined as number of seeds produced divided by maximum number of seeds that could have been produced. The homogeneity of variances is0.137. No significant different groups were found (p= 0.285). The chance for one group is 0.285 (Tukey test). 35
The highest average percentage of seed set was found with cross double pollination and self morning pollination (45% and 43%, respectively). Without pollination, a seed set percentage of 35 was found. Double cross pollination increased the percentage seed set (45% instead of 34% or 43% at single cross pollination). Afternoon cross pollination resulted in an almost as high seed set percentage as double pollination. Values do not differ significantly (ANOVA). Double self pollination was more effective than self afternoon pollination (38% instead of 26%), but morning self pollination was most effective (43%). Except for the morning treatments, cross pollination was more effective than self pollination. 36
Anthriscus sylvestris- visitors The visitor guild composition of A. sylvestris was investigated by transect observations, plot observations and verge composition.
Visitor guild The easiest way to get an idea of the visitor guild is to make transect observations. Data are presented in figure 20. Raw data are presented in Appendix M(transect A. sy!vestns)
20. Relative abundance of Anthriscus sylvestris visitors (n=352)
20
V 15 V. C 0) C 10 - 0 C) 5 C 4) I- 4) a. 0
ci. . '.. . Uj H. . I
Insect species
Figure 20. Anthrlscus sylvestris visitors. Species names of visiting individuals were recorded during transect observatIons. Insect species were grouped. The nine most abundant species are shown. The total number of visits was 352. A proportion of 0.875 is provided by the nine main visitors.
The main visitors of A. sylvestris were Empis tessellata, Eristalistenaxipertinax, Musca sp., Canthans sp., Scatophaga stercoraria,Syrphidaerest, Elateridae, Scatophaga x and Diptera small. The frequency distribution of Syrphidae-Diptera-Coleoptera-rest was 80-165-63-44. Appendix M gives also the insect species that are grouped into 'rest'. This frequency distribution can be compared to the frequency distribution on other plant species, like Aegopodium podagraria (figure22).
4 37
Plot visitor guild When plot observations are made, the time component of the visitors is also takeninto account. Plot observations are more accurate than transect observations. Becausethe large number of visitors per time unit, only small plots were used to obtain the dataneeded for figure 21. Raw data can be found in Appendix K (plot A. sylvestris).
21. Visitors on Anthriscus sylvestris
2 6 1 2 n=6 10 1 2 3 4 1
25
20 U) a) E15
0 n10 >5U) I • 0 -JRm ( 'I * •U) . I . s. . : • . Q, ( ' E •a . ! • " ' E. .! ';; ., • ..o • •d a . c • a. . t ! 0 (I, . %i LU e
Insect species
Figure 21. Plot visits (10 mm.) on Anthriscus sylvestris. Data were derived on 5 June 2001 in one verge (see AppendixA). The number of visits per insect species per umbel is given. N is the number ofobserved individuals. The number of observed insects per species is very low.
The most abundant species was E. tessellata, like in the transectobservation figure. Musca sp. is also present, but Cantharis sp. is not. Eristalis tenax/pertinaxdidalso make a large number of visits. 38
Visitor guild Aegopodium podagraria Visitor guilds of an other umbellifer in road verges, but with a later floweringperiod (Aegopodium podagraria), can be compared to the visitor guild of A. sylvestris.Raw data are presented in Appendix U (colour A. podagraria).
22. Visitors on Aegopodiumpodagraria (n = 352)
140
120 0 100 80 60 C 4.. U) > 20
0
insectspecies
Figure22. Visitors on Aegopodium podagraria. The number of visits on A. podagrarla was observed during the colourvision experiment. 364 visitswereobserved. 0.97 of the visits is represented by those nine main species.
The nine main visitors of A. podagraria were Helophilus small and Helophilus large, Enstalis arbustorum/abusivus, Melanostoma sp./P!atycheirus sp., Gymnochaetaviridis, Musca sp., Sarcophaga sp., Anasimya sp. and S. stercorana. It isremarkable, that so many half- specialised species (Syrphidae) visited the umbels. The frequency distribution of Syrphidae-Diptera-Coleoptera was 306-59-0. 39
Anthriscus sylvestris- effectiveness of visitors The flight distances and pollen loads determine among others the real value ot a pollinator for pollination. To transform transect observations to plot estimations, visitation speed and visitation density are important factors. Data are shown in figures 23 and 24. Raw data can be found in Appendix 0 (following A. sy!vestns).
Flight distances Larger flight distances between umbels mean a higher chance for transfer of pollen to genetically different plants.
23. Flight distances of Anthriscussylvestris visitors
E 0 5 2 3 3 n=2 2 4 13 6 2 12 2 0100 .0 E 80 C
60
I 40 I 20111110 il.l : E . .u).- t .0 E u a .tI; 0. Uj02 .' uJ —
Insect species
Figure 23. Average flight distances of Anthriscus sylvestris visitors. Data are derived from following observations. N is the number of followed Insects. Means plus standard errors are shown. The homogeneity of variances Is 0.287. The significance for equal means is 0.270 (ANOVA). According to the Tukey test, the significance for one group is 0.102.
Sarcophaga sp. had the largest flight distance (65 cm), followed by Cantharis sp., Empis sp. and Helophilus yellow (43, 39 and 38 cm., respectively). Tachina fabricia, E. arbustorum/abusivus, E. tenaxipertinax,Empisdiagramma, E. tessellata, Eristalis horticolaand Musca sp. are intermediate, with values between 27 and 15 cm. Empis lividahadthe smallest value of 4 cm. Differences are not statistically significant. 40
The maximum flight distances of A. sylvestris visitors are given in table 4. The maximum flight distance gives the potential efficiency of a visitor.
Table 4. Maximum flight distance between umbels by Anthriscus sylvestris visitors.
c . . E . • E Insect . .8 .. . species U) . . U) C • ' 'S... (I) . ci. ( 2 U . ' c. . .! .. (J).2 -- -. '- --- ..a - L I LU UJ LUcj Li LU. ( ()
Maximum flight 32 7.5 25 100 40 19 80 50 75 100 25 80 distance (cm)
Empis sp. and Sarcophaga sp. not just had a large average flight distance, also their maximum flight distance was largest of the visiting insect species (100 cm). The maximum flight distance of E. ilvida was only 7.5 cm. 41
Visitation speed An important characteristic of visitors is the speed with which they visit umbels. Data derived from following observations are given in figure 24.
24. Visitation speed of Anthriscus sylvestris visitors 0.30 n=2 2 4 11 8 2 12 2 5 2 4 4 c ab a ab ab ab ab ab ab b ab ab a 0.25
0.20
0.15 —.
E 0.10
C) 0.05 > 0.00 : .1 1 II t
E Uj0 . — LU Li
Insect species
Figure 24. Average visitation speed of Anthriscus sylvestris visitors. Data are derived from following observations. Speed Is defined as numberof umbellules per second. N is the number of followed insects. Means plusstandard errors are shown.
Tachina fabricia visited most umbellules per second (0.22). This value differs significantly (ANOVA) from E. livida (0.03) and Canthans sp. (0.04). 42
Behaviour Insects use their time most effectively for the plants, whenthey visit each time an umbellule of a different umbel. This increased the chance ofquickly transferring pollen to a genetically different plant. The average percentage visitedumbellules per umbel of A. sylvestris is given in figure 25.
25. Percentage visited Anthriscus sylvestris umbellules per umbel w E a. ,0 100
2 80 E
60
40 a,
C 20 U 0L. 0. 0 .. .Jil 9 I I Insect species
Figure 25. Average percentage visited umbellules perumbel of Anthriscus sylvestris. Data are derived from following. N is the numberof followed insects. Means plus standard errors are shown. After alog-transformation, the homogeneity of variances was 0.186.An ANOVA showed that the significance for equality of meansIs 0.008. The Tukey test says that there is one group with a significance of 0.14.
The average percentage of visited umbellules per umbel washighest for Cantharis sp. (85 %). Eristalis arbustorum/abusivus, E. tessellata andT. fabricia were intermediate with respectively the values of 55.9 %, 55.1 % and 55.4 %. The othervisitors had values between 18.5 % (E. livida) and 37. 5 % (E. horticola). The differences are notstatistically significant (ANOVA). 43
Pollen loads The larger the pollen load on the body of an insect, the more pollen grains can be deposited at once or the more female umbellules can be provided with pollen. If a pollen load is very small, the chance of leaving pollen to the right place on the stigmascould be very low. Raw data are presented in Appendix 0 (pollen load preparations A. sylvestris).
26. Pollen loads of Anthriscus sylvestris visitors 0 A. sylvestns pollen U rest pollen
n=13 13 4 3 3 9 10 4 6 5
9000
(0 .E 6000 I-(5 C C) 03000 Q.
0 c. '5 C..) a g. t9 . i!.. . '5 C,) UJI B L
Diptera Coleoptera
U pollen species: 7 7 6 8 7 3 3 7 3 4
Figure 26. Pollen loads of Anthriscus sylvestrls visitors. N is the number of slides. Each slide Is made of the pollen load of one visiting insect. Means plus standard errors are shown. Pollen were divided into A. sylvestris pollen and other pollen. The total number of pollen species is also given. The homogeneity of variances for the number of species was 0.165. An ANOVA could be done to compare the means. The significance was 0.002. According to the Tukey test, one subset existswith a significance of 0.059.
Eristalis sp., T. fabricia and Cantharis sp. had the largest number of pollen grains (6878, 5803 and 5656 respectively). Scatophaga x, Diptera small and Coleoptera carried the smallest number of pollen grains (1103, 407 and 909 respectively). The.other species carried between 2132 (Lepidoptera) and 3351 (E. tessellata) pollen grains. Helophilus sp. had the lowest percentage of A. sylvestris pollen grains (0.90), Cantharis sp. had the highest percentage of A. sylvestris pollen grains (0.994). The S. stercoraria and Cantharis sp. had only 3 pollen species in their loads, whereas E. livida and Lepidoptera had 9 pollen species in their loads. ______
44
Table 5 shows the fraction A. sy!vestns pollen in the loads of the visitors. Thenumber of pollen species in the loads is shown in figure 26. The pollen loads of Anthriscussylvestris visitors are very pure: all have a fraction A. sylvestris pollen above0.89.
Table 5. Fraction Anthriscus sylvestris pollen In the pollen loads of Anthriscussylvestris visitors.
Ca c I- cc d. w Insect species E C" d. . - , . U) . E . .. . 6. - E.c E E c,),< LU LU LU. Lu i-,c, o
Fraction Ant hriscus 0.9780.9680.9830.9240.8980.984 0.991 0.9650.9800.9370.9740.9950.956 sylvestris pollen
Fluorescent dye powder The results of overall activity of visitors can be shown by using fluorescent dye powder. Figures 27 a, b and c show the results. Raw data can be found in Appendix T (A.sylvestris fluorescent dye powder).
Red particles were not much found at the source (See Appendix T). In the slides they were hardly present. They seem to have a constant number atdistances of more than 5 meters. A decay to the north of the source was visible for yellow and green, and slightlyfor red. In the south, more particles were found at the same distance than in the north. In the north, up till 4 meters, there was a decay. Up till 40 meters to the south, a decay wasfound. Further away, the number of grains was higher again.
27a. Red fluorescent dye powder on Anthrlscus sylvestrls
U 0.8- 0 0 0 0 0.6 U a 0 >. 0.4 0
0.2 U II. If . e 100 -20 0 20 40 60 60 Distance from source (meters) 45
27b. Yellow fluorescent dye powder on Anthriscus sylvestris
V U 0 V S V V U S a.
0 C 0 U S U.
.20 0 20 40 60 100 Distance from source (meters)
27c. Green fluorescent dye powder on Anthriscus sylvestris
V U 0 V S V V U — - ——-—--— - S a. V V 0.4 - 0 C 0 U S U. + U • S
80 100 •20 0 stance from t°ource (meter62)
Figure 27 a, b, C. Red, yellow and green fluorescent dyeparticles on Anthriscus sylvestris. Data are derived from fluorescent dye powder (= pollenanalogue) experiment In verge 20.5. The source was at 0 m. The positive values on thex-axls are south of the source, the negative values are north of the source. After one day, at different distancesumbellules were collected. Means plusstandard errors are shown. Fluorescent dye particles seen as red, yellow and green under the UV microscope were counted in 10slides at different distances from the source at 0 m. 46
Colour vision of fluorescent dye powder Some insect species might have preferences for visitation of specificcolours. If those preferences are not evenly distributed over the insect species, thefluorescent dye powder results may be biased. Results of the colour visionexperiment on A. podagraria are given in figure 28. Raw data are presented in Appendix U (colour A. podagraria).
28. Visitors on umbels of Aegopodium podagrariawith fluorescent dye powder
n= 43 74 103.5 31 7.5 8.5 2.5 20.5 300.5 100% U) Oblancol2 C)% > 0 white part of yellow Dyellow 0 60% (II white part of orange 40% •orange fli white part of red 20% Ured
0% . U) .5 U) .tQ0 .5 .U) s. U) t'. a . Cl)
CU . Syrphida Diptera Insect species
Figure 28. RelatIve abundance In percentages of visitors oncoloured umbels of Aegopodium podagrarla. Data are derived from the colour vision experiment. White partof yellow means the white side of the umbel that was coloured yellow at one side. Thenumber of visits on the blank umbels was divided by two, because they were twice as large (hadtwice as many umbellules) as the other treatments. N is the number of visitors perInsect species. The total number of visitors on the seven treatments was calculated tobe 300.5. The percentages are given for insect species with the highestvisitation frequency. The rest Is given in rest. Also the total distribution is given.
In total, there was a slight more preference for uncoloured umbels.Most species did visit red powdered umbels, except the Sarcophaga sp.. Sarcophaga sp. andMuscasp.did both not visit the white parts of red umbels. Orange umbels were visited most bySarcophaga sp. (2/7.5). Sarcophaga sp. visited many white parts of orange umbels (3/7.5), whereasE. arbustorumlabusiVuS did not often visit this type of treatment (2/43). Yellow umbels werehardly visited by G. viridis (0/1 2.5) and Helophilus small (1/1 03.5). The white partsof yellow umbels were visited by all species except Sarcophaga sp.. Gymnochaetaviridis had a preference for blank umbels (4.5/12.5). 47
4. Conclusion and Discussion Specialisation levels Insect species can be divided in specialised (Apoidea andwasps), half -specialised (Syrphidae) and unspecialised (other Diptera). The more difficult theflower, the more specialised insects are expected. Figure 29 shows the visitor groups for fourplant species that have been described in this research. Unspecialised insects are Coleoptera and other Diptera. Syrphidae arehalf-specialised insects, Apoidea are specialised insects.
29. Fraction of insect visitors with differentspecialization levels
U specialized Dhalfspecialized • unspecialized
n=352 364 15 47
'a 0 0.75
U 0 (0 0.5 C C 0 0.25 U I-(5 U- 0 H
Plant species
Figure 29. Relative amount of insect visitors with differentspeclalisation levels. Data are derived from plot (Aegopodium podagraria) and transect(A. sylvestris and G. hederacea) observations. N is the number of visiting Individuals. Visitorsof A. sylvestris, A. podagraria, G. hederacea female and G. hederaceahermaphroditic were grouped into specialised, haif-specialised and unspeclalised visitors. Per plant species,the relative amount of these groups is given.
Anthriscus sylvestris had the most unspecialised visitors if the four taxa arecompared (0.77). The other visitors (0.23) were half -specialised. Aegopodiumpodagraria on the contrast, was visited mostly by half-specialised visitors (0.86).Unspecialised visitors did also visit A. podagraria (0.14). Specialised visitors did hardly visit A. sylvestris or A.podagraria. The visitor type composition of G. hederacea hermaphroditic resembled that of A.podagrana (0.93 half- 48 specialised), whereas the visitor type composition of G. hederacea female wastotally different. Visitors of G. hederacea female were mostly specialised (0.59), butunspecialised and half- specialised insect species did also visit this plant type (0.26 and 0.14 respectively). Umbelliferae attract half- or unspecialised visitors. G. hederacea hermaphroditic attracted haif-specialised visitors and G. hederacea female attractedmostly specialised visitors. This is strange, because specialised visitors were expected on G. hederaceahermaphroditic. The length of the whorl of G. hederacea hermaphroditic could be too largefor most specialised visitors, which have to go to G. hederacea female to drink nectar. HaIf-specialisedvisitors with a long tongue (R. campestris) can obtain pollen and nectar at G. hederaceahermaphroditic. One could argue that R. campestris is specialised if you make the subdivision on a functionaVmorphologiCal basis rather than taxonomically. HaIf-specialised visitors with a short tongue can obtain pollen and small amounts of nectar at A. sylvestns orA. podagraria.
Table 6 gives an overview of the data presented in graphs and tables on G.hederacea in the report. Glechoma hederacea- background information The experimental breeding system experiment showed that the female type had ahigher seed set than the hermaphroditic type (not significant). This agrees withWiden & Widen, 1990: seed set per fruit is higher in female than in hermaphroditic G. hederacea.Double pollination treatments resulted in higher seed set than single pollination treatments.Maybe the first deposition was not enough to trigger the enzyme system. A second possibilityis, that the first load triggered the enzymes which could be used to germinate pollen of thesecond load. Self pollination was possible, though cross pollination resulted in higher seed set. Selfpollination could result in inviable embryos. The viability of the seeds was not tested in thisresearch. The field breeding system experiment shows that for hermaphroditic plants, theavailable insect population was not sufficient to cause maximum seed set, which isfour seeds per flower. For female plants, extra hand pollination had a negative effect on seed set.The stigmas can have become too crowded or damaged by the treatment. If figure 2 and figure 3 arecombined, the female plants were visited at least one time and the hermaphroditic plants werevisited at least two times (1.9 seeds). A calculation can be made of the overall chance for seed set with selfand/or cross pollination. With this value, harmfulness of low flight distances can be computed. G. hederacea produced nectar fastest during the day. Actually, therecould be no production at all during the night, when there is no light for photosynthesis.During the night, production of nectar means allocation of sugars. The values found could be due tothe fact, that flowers were left from 16:00 the previous day, when it was still light. If theday lasts twice as long as the night, however, a decrease of nectarproduction speed of one-third overall is expected when the night is taken in. Overnight values were smaller. Maybe a touch duringthe day triggers the system to produce necessary enzymes and hence more nectar. Extra information on visitation times during the day can elucidate the foundresults on nectar production of G. hederacea. At L. album, overnight values were 5 to 10 times as low as values in thedaytime. Maybe the touch of the early insect should trigger the nectar-producing system.In that case, it could be favourable for insects to fly at little-rewarding flowers in the morning, to assurethemselves of their evening dish. Lamium album produced ten times as much nectar as G. hederacea. Overnight, concentration in the little bit of nectar was higher than in the larger amounts produced during the day. It seems, that triggering lasts for only two or three hours.'10 to 16' had a decreased concentration in comparison to '10 to12' and '10 to 14' . This could be caused by extra production of more watery nectar. '12 to 16' had also a decreasedconcentration in comparison to '12 to 14'. ______49 Tablec,rah 6a. with An both overview female of and traits hermaphroditic of visiting insect Glechoma species hederacea, on Glechoma both hederacea.values are given Data in are the directly order (female] derived (hermaphroditic].from the graphs. In the case of a C) V C) 4- . 4-0. C)C C C)C 0.n .! > C.- .0 • . 0 > C1..U).— 2 2 C) D)WC)I-> U)U = 0.0 CU 0 =)a =VO0.2g(, Lepidoptera..E In ClO 0.07;0.000.CEin.0°E4-00 9) EEc ' .w =..E>OV0.'-C) G)0ø'-. (LE 0 50-•C) 1.3Czu HelophilusEristalis sp. sp. 0.00;0.060.00;0.022i E a.E- a.E.o- h 5 18 16 RhingiaMelanostoma campestris sp./Platycheirus sp. 0.0009;0.00300.001 0;0.0038 0.00;0.470.1 3;0.1 5 18.511.0 45 3.13.5 9.185.34 25.4011.57 53.1;28.137.5;34.7'- 646 43 0.690.94 95 2.7-6.02.1-5.0 Syrphidae rest .( 2.5 ApisDiptera mellifera small 0.0086;0.0141. 0.47;0.020.07;0.09 10.2 13 13.6 9.42 10.23 54.8;44.0 59 0.47 6 3.0-6.3 BombusBombuspascuorum pratorum 0.0012;0.0083 0.27;0.17 18.9 11.82 10.89 57.6;44.2 2937 0.87 132 1 3.4 4 ApidaeBombus largesmall sp.terrestris - -______0.00;0.02 11.2 35 5.330 ?-6.4 3.32.7 2 50
Glechoma hederacea- visitors In the introduction, an estimation of insect visitors was made. Figure 31 in Appendix B gives the visitors found by Frank Hoffmann in 2000. Figure 30 compares figure 8 with figure 31. The female and hermaphroditic data of figure 8 were pooled. The areas of figure 31 were pooled.
30. Visitor guild composition Glechoma hederacea n22 62
100%
80% Lepidopfera •Enstalis sp. U Helopliussp. > 60% U Melanostoma sp./Platycheirus sp. 'I- o IJ Rhingia campestns DOiplera small • Apis mellifera 40% 0 &mbt, terresfns 0 Bombus pratorum a Bombus pascuorum • Bombus jonellus 20%
0%
Year
Figure 30. Visitor guild composition Glechoma hederacea. N is the number of observed insects in total. Data are derived from transect observations.
There was a large difference between the visitor guild composition in 2000 and 2001. In 2000, the most frequent visitor was Diptera small. In 2001 • the most frequent visitor was R. campestris.In2001, Syrphidae were found in addition to the guild of 2000.
WidenandWiden (1990) said, that visitors on G. hederacea were amongst others Lepidoptera. In one verge (verge 2) in the research area, Lepidoptera visited G. hederacea,but they can not be said to be regular or common visitors in the researched area. A visitor with a high frequency could be a bad pollinator and a visitor occurring in a low frequency could be a very good pollinator. The higher the fraction visited flowers in a plot, the more pollen will be transferred within the plot. Many flowers will receive pollen grains coming from plants in the same plot. Because G. hederacea, and especially the female form, is a donal plant, the chance is high for hermaphroditic plants that the pollen do hardly differ genetically from the receiving plant or flower. A high intra-plot visitation could enhance self-pollination. Self pollination 51 is possible for G. hederacea but the effect on the germination of the seeds and fitness of the offspring was not investigated. Pollination of female plants can only occur when a visitor deposits pollen of a previously visited hermaphroditic plant. Visitors on female flowers with a high flower constancy will almost never take pollen grains to the plants they visit. The difference in visitors on the plant forms at the transect observations can be caused by the difference in food necessarity of the visitors. R. campestris eats pollen grains and drinks nectar, which results in the choice of hermaphroditic flowers, which provide both sources. A. mellifera has a shorter tongue than R. campestris.A.mellifera uses nectar as energy source and is able to take it out of the female flowers, which have a shorter calyx. A. mellifera can also be a collector, but individuals with corbiculars filled with G. hederacea pollen were rarely observed. The proportion of Lepidoptera is based on the observation of one Lepidoptera species in a small population of female G. hederacea plants. Difference in Lepidoptera between the plant forms is biased by incidence. The difference in visitors from the methods plot observations and transect observations is obvious. The role of B. pascuorumonhermaphroditic flowers is in figure 8 (transect observations) not as distinct as in figure 7 (plot observations). The difference in number of visits on the plant types is only apparent in figure 7. Differences can be caused by the inaccurate observations of transect walks: the duration of visits is not taken into account. When a visitor is very slow, it can be spotted during transect walks. There is a large chance that the slow visitor does not reach the plot, which is only observed during ten minutes. The larger the plot, the more complete view results about the visits made by the visitors. Figure 11 shows the visitation speed of G. hederacea visitors. Those values could be used for the transformation of transect observations to plot estimations. Because no information about visitation speed is known from insects only observed in transects, this calculation can not be done. A fast visiting insect has a higher chance to be seen in a plot observation of 10 minutes than a slowly moving insect. If this theory would be true, faster insects score better in plots than in transects. Apis mellifera and B. pascuorum were faster than Melanostoma spiPlatycheirus sp. and R. campestris. On hermaphroditic flowers, indeed Apis mellifera scores better in plots than in transects. This is partly true for B. pascuorum. On female flowers on the other hand, this can not be seen. Visitors on G. hederacea plants have the same visitor rate in the three areas. Glechoma hederacea- effectiveness of visitors Statistically, there is no difference in foraging distances. The larger the flight distance, the higher the chance for mixing of pollen with non-siblings. Rhingia campestris will take pollen over large distances and so enhance cross-pollination. Bombus sp., Melanostoma sp.IPlatycheirus sp. and A. mellifera did not take pollen over large distances and could enhance inbreeding by taking pollen to siblings. The inbreeding chance probability at a certain distance will differ between populations, because every population has a different genetical structure. The higher the visitation speed, the more flowers per time unit are visited. If a visit takes long, extra pollen could be deposited by the movements of the visitor. Bombus sp. and A. mellifera visited many flowers per time unit. Melanostoma sp./Platycheirussp.and R. campestris visit less flowers per time unit. The importance of Melanostoma sp./Platycheirus sp. and R. campestris could be underestimated by transect observations. Depending on the size and visiting behaviour, each insect species will have its own visitation speed and pollen deposition. This should be investigated further to make a good conclusion about the residence times found. Bombus sp. was the fastest visitor (inflorescences per minute), followed by A. mellifera. Melanostoma sp./Platycheirus sp. and R. campestris are slower visitors. For bumblebees, a foraging rate of 10-20 flowers a minute is common (Heinrich, 1979). This is in agreement to the value 18.9 flowers per minute. The number of visited inflorescences per minute could be a better measure for the visitation speed than the number of visited flowers per minute, because when a new inflorescence is visited, the flowers get foreign pollen. However, self-pollination is not impossible in G. hederacea (figure 2). The values of R. campestris are strange: four times as many flowers as inflorescences are visited per minute, and still the visitation speed on inflorescences is three times as high as that on flowers. The visitation speed on inflorescences was expected to be four 52 times as low as that of flowers. This means, that the followed individuals in plot observations were twelve times as fast as the followed individuals during following observations. It is possible that the followed individuals were extraordinary slow. The plot observations are more precise: they were done in different verges. Rhingia campestris and Melanostoma spJP!atycheirus sp. may enhance cross-pollination more than A. mellifera and Bombus pascuorum do. The differences are not significant. Glechoma hederacea is self-compatible (figure 2), but the fitness of the population will stay higher when foreign pollen grains are received. The higher the fraction of visited flowers from presented flowers, the less cross- pollination will occur on hermaphroditic flowers: only changes from one inflorescence to another of a different clone could cause cross-pollination. Insect species with a foraging behaviour that results in lower flowers of inflorescence visited fractions are said to be potentially better pollinators. Of course, also their flight distances should be taken into account. Rhingia campestris seems to be the least effective pollinating visitor in this view. Not only the load is important, also the deposition. Unfortunately, no research has been done to the optimum number of pollen grains deposited for pollination. Half of the bumblebees carry two or more kinds of pollen (Heinrich 1979). Only 2% of the honey bees carry 2 kinds of pollen (Heinrich, 1979). Apis mellifera was found to have 3 to 6.3 pollen species on average in its load, and B. pascuorum 3.4. The values of Heinrich about A. mellifera do not agree with the values of this research. In my investigation A. mellifera was less flower constant. Better pollinators leave pollen grains of only one or two species. Their foraging behaviour is flower constant. The most flower constant visitor were Lepidoptera, the least flower constant visitor was B. pratorum. On the first pistil, more pollen grain species are expected than on next pistils, because the insects get rid of their pollen loads during their visits. An explanation for an increase of pollen grain species on next pistils is, that the pollen loads are built up of several layers. If the upper layer of pollen species A is left on a pistil, the next layer can be left on the next pistil. In contrast to data from pollen loads, those data give insight in effective flower constancy (layer theory). Further research should be done on the pollen loads of Lepidoptera. The little abundant group seems to have interesting pollen load deposition numbers on G. hederacea. Fluorescent dye particles can be transferred over 64 meters in two days. Decay is not obvious however. There might be so many traffic, that all flowers over a large area have almost the same number of dye particles, the number depending on how attractive the flower is. The source could be set at 100%. The radius of a foraging bumblebee is found to be maximally 450 meters in certain circumstances (Kwak & Tieleman, 1 994c). Herefore, we could expect fluorescent dye powder to come maximally 450 meters, if the load is not depleted beforehand and if no other insects transfer the powder further. Transferred dye powder can be transferred from one stigma to the next. The same process could be at work for pollen grains, but because of size differences, this process can be enhanced with fluorescent dye particles. There could be a difference in adhesion between fluorescent dye particles and pollen grains. Fluorescent dye particles are lighter, and might be more sensible for wind. The estimated mean pollen dispersal of G. hederacea is 5.3-5.9 meters (Widen & Widen, 1990, data from seed set observations). The value of 64 is eleven times as high, but is a total of added means and carry-overs. Glechoma hederacea- important pollinators Here, I propose to combine traits to come to a conclusion about pollinator importance. I will like to assign the insect species most favourable for pollination a value of 1, and insect species less favourable for pollination a proportional value. Because no extra information is available, I assume that the values of the traits are correlated with favourability linearly. I assume equal importance of the traits for effectiveness, though opinions on this case differ. A value lower than 1, e.g. 0.5 for trait A. means that the insect species will pollinate 0.5 times as many flowers as the insect species with the value of 1. A value of 0.25 means that the insect species will pollinate 0.25/0.5=0.5 times as many flowers as an insect species with value 0.5. If the insect species with value 0.5 for trait A. has a value of 0.5 for trait B., the insect species will be able to pollinate 0.5 x 0.5 = 0.25 of the flowers that an imaginative insect species with best values for all the traits could have pollinated. The values for the traits of one insect species have to be ______
53 multiplied to obtain a total pollination importance measure. Only the (imaginative) insect species that combines the best values for all the traits has an effectiveness of 1. The chosen traits for calculation of effectiveness of visitors of G. hederacea are: 1. proportional number of observed visitors by transect observations and number of visited inflorescences per second: transect observations times number of inflorescences per second gives more accurate information about the pool and the number of inflorescences visited by this pool than plot observations. 2.number of inflorescences visited per minute: only flights from iriflorescence to inflorescence are supposed to be effective 3.flight distances: a measure for the inbreeding chance. 4.species specific pollen deposition: gives more information than pollen load. The proportional number of observed visitors is split up into female and hermaphroditic G. hederacea. Therefore, the total effectiveness on both female and hermaphroditic G. hederacea can be calculated.
Table7a. An overview of indexed effectiveness values for Glechoma hederacea female visitors. In the case of missing values, averages were taken (italic script). Insects with too many missing values have been left.
0C C u .3 0 W CI)C .2c U) . C.) o •—Q O) c c. (.)0 U) ..- Co 0. 00 t. .° '-00.U). — U) .0U)O° 0t.!i.. U .c U)U) U) E U) co 0 . Z.9? I Z.5. Melanostoma 0.28 0.59 0.45 0.59 0.044 spiPlatycheirus sp. Rhingia campestris 0 1 0.78 0.35 0.000 Syrphidae rest 0.62 0.71 0.76 1 0.334 Apis mellifera 1 0.55 0.80 0.22 0.097
Bombuspascuorum 0.57 0.71 1 0.48 0.194
Table 7b. An overview of indexed effectiveness values for Glechoma hederacea hermaphroditic visitors. In the case of missing values, averages were taken (italic script), Insects with too many missina values have been left.
C . 0 C.) U3C U) co— CØ U) o t 5 3 E '2 0 . co.C .) cD 13 C_ QC w .c- woE0 -Q).5 OEU)U) Z.c LI.. Z.c5( o. i-._.c.c Melanostoma sp./Platycheirus sp. 0.32 0.59 0.45 0.59 0.05
Rhingia campestris 1 1 0.78 0.35 0.273
Syrphidae rest 0.19 0.71 0.76 1 0.103 Apis mellifera 0.04 0.55 0.80 0.22 0.004 Bombus pascuorum 0.36 0.71 1 0.48 0.123 54
With this method, Syrphidae rest and B. pascuorum appear to be the most important visitors for pollination, on female G. hederacea. On hermaphroditic G. hederacea, R. campestns and B. pascuorum are the most important visitors for pollination. If those species would become extinct, a lot of pollination would not be done any more, but the gap may be filled by other visiting insect species. In figure 30 was shown that the visitor guild composition can have large differences between years. The pollination importance of the visiting insect species in 2001 differed probably from the pollination importance of the visiting insect species in 2000.
Table 8. An overview of traits of visiting Insect species on Anthriscus sylvestris. Data are directly derived from the graphs. 4- I- U) .c 0 2 2 w 0. U 0 2d U) 0 0 U) W '4 0. ' V U) U) U) ° S LE 4- ..-c g 0 4- .CU) Ec 2 c c —U) 0 o •.o>E2 Th >EE .E o*LLD *5 — ua Lepidoptera 0.97 Empis livida 7.5 0.03 19 0.96 Empis diagramma 6 21.3 32 0.11 27 0.98 Empis tessellata 19.3 21 16.6 25 0.11 55 3293 0.98 Empis sp. 39.1 100 0.11 24 Eristalis arbustorum 7 26.5 40 0.16 56 Eristalis horticola 4 15.4 19 0.18 37 Eristalis tenax/pertinax 17.9 8 23.5 80 0.17 31 6355 Eristalis sp. 0.92 Helophilus trivittatus 50 0.12 26 0.90
Rhingia campestris 1 Syrphidae rest 4.8 2952 Tachina fabricia 3 27.2 75 0.22 55 5708 0.98 Sarcophaga sp. 5 65 100 0.16 28 Scat ophaga stercoraria 6.2 2344 0.99 Scatophagax 3.7 1065 0.97 Musca sp. 13.9 11 16 25 0.07 34 2377 0.98 Diptera small 3.7 1 381 0.94 Cantharis sp. 13.9 2 43.8 80 0.04 85 5626 0.99 Elateridae 4.0 Coleoptera rest 869 0.96
Table 6b gives an overview of the data presented in graphs and tables on A. sylvestris in the report. Anthriscus sylvestris- background information In the studied literature, no information was available about the breeding system of the species A. sylvestris. From the breeding system experiment can be concluded that A. sylvestris is highly self-compatible. Cross pollination might result in a higher seed set percentage because of the lack of inbreeding depression, but the difference is not significant. As in G. hederacea, the lowest seed set was caused by the treatment of self pollination at 3 p.m. The experiments in the garden were done with netted plants. The netting of those plants might have been not as adequate as needed. This can have caused insects to pollinate flowers or intrude under the nets. Next time, some reference preparations should be made to control the virginity of the stigmas. A problem with A. sylvestris was, that the wind could have blown the 55 plantsto each other and hence cause pollination. A netting method of small plant parts instead of a group of plants is recommended. Anthriscus sylvestris- visitors Unspecialised visitors like other Diptera and Coleoptera were expected. Diptera and Coleoptera are two-third of the total visitors. Other Diptera and Coleoptera are unspecialised species. Still, almost one quarter of the visitors are half-specialised insects: Syrphidae. The difference in visitors from the methods plot observations and transect observations is obvious. Differences can be caused by the inaccurate observations of transect walks: the duration of visits is not taken into account. When a visitor is very slow, it can be spotted during transect walks. There is a large chance that the slow visitor does not reach the plot, which is only observed during ten minutes. The larger the plot, the more complete view results about the visits made by the visitors. Figure 24 showed the visitation speed of A. sylvestris visitors. Those values can be used for the transformation of transect observations to plot estimations: Eristalis tenaxipertinaxhasa high visitation speed. Still, its value is lower in plot observations than in transect observations. The same is true for Muscasp.The theory does not fit to these data, but the difference between the methods is obvious. Although unspecialised visitors were expected, haif-specialised insects visited the umbels of A. podagraria. The visitor guild was totally different from the visitor guild of A. sylvestris. Both plant species are very common Umbelliferae but they differ in flowering time. Anthriscus sylvestris- eftectiveness of visitors A larger flight distance is better for pollination: there is a higher chance for mixing different gene pools. Sarcophaga sp. was the most favourable visitor, whereas E. livida was not of much help at pollination in this view. The lower the visitation speed, the more time is lost, that could have been used for pollination. Empis livida and Cantharis sp. were significantly slower visitors than T. fabricia. Visitors with a very high speed can be worse pollinators, because there is not much contact between the animal and the stigmas, resulting in less pollen deposition. The lower the percentage visited umbetlules per umbel, the more effective the behaviour is for cross-pollination. Empis livida, Empis sp., E. diagramma, Helophilus yellow, E. tenax/pertinax,Sarcophagasp., Musca sp. and E. horticolaweremost favourable visitors for pollination in this view. Eristalis arbustorum/abusivus, E. tessellata and T. fabricia were intermediate, whereas Cantharis sp. was least favourable for pollination. The Diptera small, Coleoptera and Scatophaga x carried the lowest number of pollen grains, and were therefore the worst pollinators. Still, on average the small fly carried 407 pollen, which could be enough for pollination. Eristalis sp., T. fabricia and Cant hanssp.carried the highest number of pollen grains, and were therefore the best pollinators in this perspective. Insect species with more homogeneous loads are more flower constant. The pollen of A. sylvestris are so small, that transport or falling down by the wind is very well possible. During the creation of slides, pollen can have reached the gel. Next time, from time to time reference slides of the gel have to be made, to be sure of a clean gel. The purity of the pollen loads of A. sylvestris stigmas could be due to counting errors: estimations of large numbers can hide other pollen species. The fluorescent dye powder experiment showed larger amounts of dye particles in the south than in the north. Maybe smaller particles were counted, or the vegetation structure in the verge was responsible for this strange outcome. Transportation by wind, either directly to the powder or indirectly to the insects, is also a possibility. At 100 meters, fluorescent particles can still be found. In one day, particles and or pollen can be transferred over this distance. Standard deviations are rather large, so the graph has no prospecting value. The decrease of the total number of fluorescent dye grains seems to be more smooth than the decrease of every colour separately. The decay of the fluorescent powder of A. sylvestris differed a little from the decay curve of G. hederacea. On A. sylvestris, the values were lower at small distances from the source than ______
56 on G. hederacea. Further away, a decay could be seen, whereas thelevel stayed almost the same on G. hederacea stigmas that were further away. For the species that visit both A. podagraria and A. sylvestris, we can draw conclusions about colour vision. Overlapping species were E. arbustorum/abusivus, Sarcophaga sp. and Musca sp. Application of fluorescent dye powder seems to have an effect on visitation of insects. However, this effect differs for different insect species. in total there is a preference for blank umbels. Both yellow and white parts of yellow umbels seem to be of no preference.
Anthriscus sylvestris- important pollinators It is a pity that pollen deposition on A. sylvestris could not be measured. For the effectiveness of A. sylvestris visitors, I will use the same method as described for G. hederacea. The used traits are: 1. relative abundance (from transect observations), which together with the number of umbellules visited per second gives the total number of visited umbellules per second by one insect species. 2.flight distance: a measure for the inbreeding chance 3. species specific (A. sylvestris) pollen in the load. I expect no negative effect on seed set of foreign pollen because of their low frequencies. Insects with too little values have been left out, otherwise averages were taken (italic script).
Table 9. An overview of indexed effectiveness values for Anthriscus sylvestris visitors. In the case of missing values, averages were taken (italic script). Insects with too many missing values have been left out.
C) C) C . C V C) cc cø C) UI V o .2c c i.Q)C) C) 0. . 50. C) O C.) c . CC) C) .o c C) ..,0 0. . .5C UI — > V 0. . .0 C . D C) .. C) E.o° (jO U) 2 .— o .E cr zc, u
Empis tessellata 1 0.65 0.38 0.52 0.128 Eristalis tenax/pertinax 0.93 1 0.54 1 0.502 Syrphidae rest 0.25 0.58 0.57 0.46 0.038 Scatophaga stercoraria 0.32 0.58 0.57 0.37 0.039 Scatophaga x 0.19 0.58 0.57 0.17 0.011 Musca sp. 0.72 0.41 0.37 0.37 0.040 Diptera small 0.19 0.58 0.57 0.06 0.004 Cantharis sp. 0.72 0.24 1 0.89 0.154 Elateridae 0.21 0.58 0.57 0.48 0.033
Enstalis tenaxipertinax is in this method the most important pollinator on A. sylvestris. Empis tessellata and Canthans sp. were also important pollinators. If those species become extinct, a pollination gap would appear, but this gap could be filled by a large range of other visitors.
General The composition of the flowering plant species differed much between verges and during the season. The variation in visitors was so high that comparison between areas was difficult. The importance of flight distance can only be calculated when the average clone sizes of the plant species and maybe even the geneticai community structure are known. Because both (3. hederacea and A. sylvestris are self-compatible, this information is not so important, because a pollen grain can germinate on any plant of the own species. Questions aboutimportance of 57 insects for reproduction can better be answered on species that do notreproduce clonally. The problems a clonal species has to cope with (fitness) are of anotherlevel than the problems of a strictly sexual reproducer (survival, then fitness!). It would be interesting to know, what the number of pollen produced perplant is: when the total number of plants stays the same, at least one of theproduced pollen should pollinate another plant. Also, the total obtainable number of pollen per visit can becalculated. The fluorescent dye powder experiment was done only once forboth plant species. If a pilot experiment had been done, we would have searched for a longer vergeto make the tail of the graph more complete. The fluorescent dye powder experiment wasnot safe, because the former colour white consisted of green and orange particles. Next time,fluorescent dye powder should be checked on purity before use. Another problem of the fluorescentdye powder was the size of the particles. Some of them were really much smaller thanothers, and hardly visible. In a view-screen with maybe 100 particles, it is difficult to estimate numbersof unequally sized particles. Therefore, a colour should be used with equally sized particles, orthe particles should be sieved somehow. Another possibility is to only count particleslarger than a certain minimal particle size. The interesting question about pollinator importance can only be solved when many information is available on both the traits of visiting insects and the importanceof those traits for pollination. To cope with questions about biodiversity changes, experiments are necessary.Some plant species can be removed and what happens to the visitor guilds ofother plant species can be investigated. It would be interesting to see if an insect community as awhole can change its preferences and visit the introduced plant species. 58
5. Acknowledgements I thank Manja Kwak and Frank Hotfmann for their help during my subject. They gave me freedom to execute my experiments and corrected me where necessary. They spent much time talking with me about the research. I'm thankful for the drives that Manja gave me with the rented bike in the back. I won't forget the hospitality after a hard day's work. I thank Jelte van Andel for his help in finding this subject for me and in doing some field work. I want to thank my parents for the motivational support I needed and some good ideas of my father.
6. References
Beaftie, A.J.,1972.A technique for the study of insect-borne pollen. Pan-Pacific Entomologist 47:82 Corbet, S.H., 1999. Spatiotemporal patterns in the flowering of bluebell, Hyacinthoides non-scnpta(l-lyacinthaceae).Flora 194, 345-356 Dramstad, W.E., 1996. Do bumblebees (Hymenoptera: Apidae) Really Forage Close to Their Nests? Jn. of Ins. Beh. 9 (2): 163-182 Ellis, W.N. and Ellis-Adam, A.C., 1994. Do ene Schermbloem is de andere niet (Umbels are not alike) Ent. Bar., Amst. 54(10): 191-199 Grace, J. and Nelsen, M., 1981. Insects and their pollen loads at a hybrid Heracleum site. The New Phyt. 87, 413-423 Helnrlch, B., 1979. Bumblebee economics — Harvard University Press, Cambridge, Massachusetts, and London, England Hen-era, CM., 1987. Components of pollinator 'quality': comparative analysis of a diverse insect assemblage. Oikos 50:79-90 Koul, P., Sharma, N. and Koul, A.K,, 1993. Pollination biology of Apiaceae. Current Science, 65(3): 219-222 Kwak, MM., 1994a. Populatie structuur & bostuiving: effecten van ruimtelijke rangschlkking bij de Zwartblauwe Rapunzel. Landschap 11(1): 15-24 Kwak, M.M., 1 994b. Planten en bestuivors: achteruitgang leidt tot verschuivonde relaties. Landschap 11(1): 29-39 Kwak, M.M. and Tieleman, I., 1 994c. Het Hommelleven — Stichting Jeugdbondsuitgeverij en Stichting Uitgeverij Koninklijke Nederlandso Natuurhistorischo Voreniging, Utrecht Kwak, MM. and velterop, 0., 1997. Flower visitation by generalists and specialists: analysis of pollinator quality. Proc. Exper. & AyI. EntomoJ. 8:85-89 Kwak, M.M., Velterop, 0. and Van Andel, J., 1998. Pollen and gene flow in fragmented habitats. App!. Veg. Sd. 1:37- 54 Memmett, J., 1999. The structure of a plant-pollinator food web. Ecology Letters 2: 276-280 Olsen, K.M., 1997. Pollination effectiveness and pollinator importance in a population of Hetherotheca subaxillaris (Asteraceae). Qecologia 109:114-121 Prins, A.H., Dijkstra, G.A. & Bekker, R.M., 1998. Feasibility of target communities in a dutch brook valley system. Acta bet. Neon. 47(1): 71-88 Rlcclardelll d'Aibore, G.C., 1986. Les insectes pollinisateurs de quelques ombellifbres d'intérét agricole et condimentaire (Angelica archangelica L, Carum carvi L, Petroselinum cnspum A.W. Hill., Apium graveolens L., Pimpinella anisum L., Daucus carota L., Foeniculum vulgare Miller v. azoncum Thell.). Apidologie 17(2): 107-124 Van der Goot, V.S., 1989. ZweeMiegen — KNNV, Jeugdbondsuitgeverij Van der Goot, V.S., 1990. Dansvliegen determineertabe! voor de wat grotere soorten van het geslacht Empis en aile soorten van hot geslacht Hybos in de Benelux - Jeugdbondsuitgevenj Van der Meljden, R., 1996. Heukels' Flora van Nedenland— Wolters-Noordhoff, Groningen, 22e druk Van Veen, M. and Zeegers, Th., 1993. lnsecten Basis Boek—Jeugdbondsuitgevenj, twoede druk Weeda, E.J., Westra, R., Westra, Ch. and Westra, T., 1987. Nedenlandse Oecologische FLORA -wilde planten en hun relaties 2. Eon uitgave van hot IVN in samonwerking met de VARA en do VEWIN; 242-290 Weeda, E.J., Westra, R., Westra, Ch. and Westra, T., 1988. Nedenlandse Oeco!ogische FLORA -wilde planten en hun relaties 3. Eon uitgave van hot IVN in samenwerking met do VARA en do VEWIN; 145-184 Widen, B. and Widen, N., 1990. Pollen limitation and distance-dependent fecundity in females of the donal gynodloecious herb Glechoma hederacea(Lamiaceae). Oecologia83: 191-196 ______Plant species veraesResearch in whichcalendar. research This tableis done gives are an in overviewthe cells. of the investigations. Different research elements are in columns, dates in rows. Glechoma hederacea Anthriscus syivestris The DatePart: ci, U) Cu C .2 C .2C C U) CU) (yyyymmdd) + W .-o0c .0wU) 0C V 0 C U) o.I- I- .0 COW .2i, V 0 c 20010502 >02,E' 5,o 9,o • .2Q0 2, 5,CU)U)Q 9, .0 = U) .5_c D0 .u 0) Z .z—. .0 _O.0C U) .c (0 0 •: 0 (/) 17.5,12, 17, O12, 17, . 0. O 0.0 25.5,24,18.5,18, 25, 25,26 2001050820010503 17,26,17.5, 26.5 18, 19.517 19.518.5,17, 18, 19.5 18.519.5 17 20010509 2, 4019.518.5,18, 2, 5, 40 2, 40,19.518, 5 2 2 2 20010510 25,26.5,26,25.5, 27.5,27,26.5,25, 26, 26.5,27.5,34,26, 27, 34 20010515 27.5,3427, 25+25.5 2 x x 20010529200105232001052120010516 5 40 2 18 20.5n, 5,36 20.536 20.5 200106122001060520010530 18 34 35 3520.5s 36 m x? L = Lamium album s=southn=north x=experimentaim=Manja's garden garden .Q CD Appendix B 31. Verge visitor composition 2000
insect species visiting Glechoma hederacea N.S N.15 N.1 100% •oth., OIsru 90% ORh4ig. canw.IU 00%
10% •Bombue jon.Iu. S. 50% BBcmbrje pre(onim
50% •Bomb,a t.n*IOst
40% H I OBc,,,bra pescua.Um 30% •Bombu, troao.'um 20%
10%
0% —J
Aria
Insect species visiting Anthriscus sylvestris
N N.110 N.ISO
100% •Muscsiae C S.
90% • Scatopfla0ae
60% •Apsdae rest
70% DH1flefl0p(l rest 60% DCcIsopstla & 50% rest 40% S •Syvplitdas 30% • BorflbuS & PSySstus 20%
10% 0j__
Figure 31. Visiting insect species composition of G. hederaceaand A. sylvestris in the three research areas. Data for G. hederacea were collected in May and June 2000 In all verges where G. hederacea occurred. Data for A. syl vest ris were collected in the fourth week of May 2000 in 9 selected verges (3 per area). (Hoffmann, unpublisheddata).
For G. hederacea, Stroomdal Drentse Aa had totally different visitation than Bovensmilde and Ekehaar. This is due to only one observed insect. The visitation on A. syl vest ris can be compared more easily, though Drentse Aa had less Apidae rest and Scatophagidae visitors than the other two areas. Appendix C 32. Nectar production by Glechoma hederacea plant types
In contrast to figure 4, this figure shows the nectar production perhour by G. hederacea, divided into female and hermaphroditic plant type.
32. Nectarproduction by Glechomahederacea, dividedIntoplant types
0 female • hermaphroditic 1221 420 n= 0 26 10 22 0 8 0 0 047 2241 656 326 0.020 C) 0 . 0.016 E I- 2 0.012
0.008
•0 0 0.004
U C) Z 0.000
Treatment
Figure 32. Nectar production by Glechoma hederacea, divided intoplant types. Treatment 10 means 10 a.m., 12 means noon, 14 means 2 p.m., 16 means4 p.m.. Nectar was depleted at 4 p.m. the day before the nectar amount measurements.There are ten 'treatments', in which the production periods and production times varied. Y errorbars are standard errors from the means. N is the number ofinvestigated flowers. The homogeneity of variances is 0.000. The chance that the means come from the samepopulation is 0.071 (ANOVA). The means can be classified in one group (Tukey) with a probabilityof 0.721.
There was no significant difference between production by female andhermaphroditic G. hederacea flowers. Appendix D 33. Pollen grains on Glechoma hederacea virgin pistils Contrary to figure 16, this figure shows the number of deposited pollen grains on the first, second and third stigmas.
33. Pollen grains on Glechoma hederaceavirgin pistils
•Pistill U Pistil 2 OPistiI3
11 2 13139111 332111 31 331 In=3 3 8 5 0 T.i U) •U) U)E E U) 1: II H —I u'.
Insect species
Figure 33. Pollen grains on Glechoma hederacea virgin pistils. Means plus standard errors are shown. N is the number of stigmas.
Because no extra pollen grains can be taken up from the emasculated flowers and pollen grains are lost during the first visit, the number of deposited pollen is expected to be highest at the first stigma. The number of deposited pollen was not highest at the first stigma. Both female and emasculated hermaphroditic plants were used. The error bars show, that the variation of deposition is very large. Appendix E Glechomahederaceabreeding system experimental hand pollination was done in the experimental garden
Clone Mother plant ILegend IPlant type I I 19-1 1 1 not Hermaphroditic 2 self 9:OOh Hermaphroditic 19-3 2 3 self 15:OOh Hermaphroditic 13-3 3 4 4 self 9:OOh and 15:OOh Hermaphroditic 19-2 5 5 cross 9:OOh Hermaphroditic 18-2 6 cross 15:OOh Female 13-2 6 7 cross 9:OOh and 15:OOh Female 18-1 7 Female 18-3 8
I Mother plant I Treatment numberiNumber iiiiiiit numberl Number of seeds I 6 1 1 0 3 6 0 1 1 0 4 6 0 1 1 4 6 2 1 0 4 6 1 2 1 0 5 6 1 2 1 5 6 0 3 1 5 1 7 2 3 1 0 1 7 3 3 1 0 4 4 1 0 7 7 2 4 1 2 1 4 1 2 7 7 3 5 1 0 2 7 0 5 1 0 3 7 2 5 1 3 7 0 1 2 0 3 7 0 1 2 4 4 7 4 1 2 4 4 2 2 0 4 7 2 2 0 5 7 2 2 2 5 7 3 3 2 0 5 7 1 4 3 2 0 6 1 3 2 6 1 1 4 2 0 6 1 1 4 2 3 7 1 0 4 2 7 1 0 5 2 3 7 1 0 5 2 0 8 1 0 5 2 2 8 1 0 1 0 1 3 4 8 1 0 1 3 7 1 0 1 3 7 2 3 7 1 0 2 3 7 1 0 2 3 7 1 0 3 3 0 7 1 0 3 3 0 6 2 1 3 3 0 6 2 0 4 3 2 6 2 1 4 3 0 7 2 0 4 3 7 2 3 5 3 2 7 2 1 5 3 0 8 2 0 5 3 8 2 0
1 4 2 8 2 0
1 4 0 7 2 2 3 1 4 7 2 2 4 1 7 2 2 4 2 7 2 2 3 4 0 7 2 4 3 4 0 7 2 0 3 4 0 6 3 2 4 4 2 6 3 3 4 4 3 6 3 3 4 4 7 3 4 4 5 4 4 7 3 5 4 2 7 3 1 5 4 3 8 3 0 3 1 1 5 0 8 0 1 5 0 8 3 3 1 1 5 0 7 2 5 3 7 3 1 2 5 0 7 3 4 2 5 2 7 3 0 2 3 5 0 7 3 0 3 5 1 7 3 3 5 0 6 4 1 4 5 0 6 4 0 4 5 0 6 4 3 4 5 7 4 2 5 5 3 7 4 4 5 5 4 7 4 1 5 5 4 8 4 4 4 0 1 6 2 8 4 2 1 6 4 8 4 4 1 6 0 7 4 1 2 6 1 7 2 6 0 7 4 0 2 6 7 4 4 3 6 4 7 4 2 4 1 3 6 1 7 Appendix F breeding system Glechoma hederacea field
Experiment in verge 34 Pollinated at 10 05 2001 Collected at 30 05 2001
Verge Plant type # seeds # seeds extra pollinationcontrol 34 hermaphroditic 4 4 34 hermaphroditic 3 4 34 hermaphroditic 1 1 34 hermaphroditic 3 0 34 hermaphroditic 2 0 34 hermaphroditic 3 4 34 hermaphroditic 2 3 34 hermaphroditic 4 4 34 hermaphroditic 2 2 34 hermaphroditic 4 1 34 hermaphroditic 4 3 34 hermaphroditic 0 2 34 female 1 1 34 female 4 0 34 female 0 2 34 female 2 3 34 female 0 2 34 female 0 2 34 female 1 3 34 female 1 2 34 female 1 1 34 female 3 4 34 female 1 0 Appendix G Anthriscus sylvestris breeding system experimental hand pollination was done in the experimental garden population came from Manjas garden, Assen a fruit can contain 2 seeds
# fruits realised % realised of possible fruits I Mother I Treatment# treated umbellules I # possible fruits I 5 38.5 blue 1 6 13 31.5 54.3 brown 1 11 58 1 16.7 darkgreen 1 3 6 9.5 52.8 yellow 1 6 18 4 19.0 light green 1 9 21 1.5 6.3 red 1 7 24 19 57.6 white 1 11 33 blue 2 9 40 26 65.0 25.0 brown 2 7 20 5 dark green 2 5 7 2.5 35.7 yellow 2 5 14 7.5 53.6 light green 2 10 29 13 448 57.1 red 2 7 28 16 white 2 8 24 5.5 22.9 43.8 blue 3 5 8 3.5 21.4 brown 3 4 7 1.5 dark green 3 8 23 6.5 28.3 yellow 3 9 42 15 35.7 18.5 light green 3 10 27 5 13.3 red 3 5 15 2 white 3 10 34 6.5 19.1 15.0 blue 4 9 30 4.5 48.1 brown 4 7 26 12.5 dark green 4 4 9 5 55.6 yellow 4 10 44 25.5 58.0 light green 4 9 26 7.5 28.8 19.0 red 4 7 21 4 white 4 12 38 16 42.1 45.5 blue 5 5 11 5 brown 5 6 19 4 21.1 dark green 5 7 29 9 31.0 yellow 5 4 12 5.5 45.8 light green 5 7 21 7 33.3 41.7 red 5 8 24 10 21.2 white 5 13 52 11 50.0 blue 6 5 15 7.5 18.4 brown 6 5 19 3.5 dark green 6 9 28 15.5 55.4 yellow 6 9 39 22.5 57.7 light green 6 10 30 9.5 31.7 51.6 red 6 7 31 16 white 6 12 35 11.5 32.9 blue 7 7 22 7 31.8 brown 7 7 31 10 32.3 darkgreen 7 7 13 5 38.5 yellow 7 4 13 6 46.2 light green 7 7 25 17.5 70.0 red 7 7 21 6.5 31.0 white 7 9 32 20 62.5 pot mt.flowertimetreat-mm. microliter nectar plant type nectar conc. 1 =temaie Appendix H 2=hermaphroditic - - -- 1 0.0313 1 13.11 1 1 1bj 10 nectar Glechoma hederacea 1 13.21 1 1 12:16 1 0 0.0000 1 13.21 1 1 16:25 5 0 0.0000 1 13.21 1 1 10:17 7 0 0.0000 pot numbers e.g.: 13.21: 13=mother verge 13.31 1 1 12:08 1 0 0.0000 2 2 = site in mother verge 13.31 1 1 16:19 5 1 0.0313 2 1= pot 13.31 1 1 10:09 7 1 0.0313 2 2 13.32 1 1 16:38 3 7 0.2188 plants had been netted in the experimental garden 13.32 1 1 10:52 7 0 0.0000 2 18.11 1 1 11:59 1 3 0.0938 46 1 from 1405 2001 16:00 plants in greenhouse 18.11 1 1 16:13 5 0 0.0000 1 0.1563 1 18.11 1 1 10:01 7 5 date of measurements: 1505 2001 1 18.12 1 1 12:35 1 0 0.0000 1 18.12 1 1 16:39 5 3 0.0938 * in 1 microliter capillary, 32 mm 1 18.12 1 1 10:58 7 1 0.0313 2 19.31 1 1 16:00 3 3 0.0938 conc. = concentration; inf.=inflorescence 19.31 1 1 9:44 7 0 0.0000 2 1 13.11 2 1 12:23 1 0 0.0000 Treatment legend 13.11 2 1 16:33 5 0 0.0000 1 1 lOtol2 13.11 2 1 10:29 7 0 0.0000 1 1 2 lOtol4 13.21 2 1 16:29 10 1 0.0313 3 lOtol6 13.31 2 1 16:21 10 1 0.0313 2 4 12to14 13.322 1 16:38 10 0 0.0000 2 5 12to16 18.11 2 1 16:14 10 2 0.0625 1 6 14to16 18.122 1 16:40 10 0 0.0000 1 7 16 to 10 19.31 2 1 11:47 2 1 0.0313 2 8 16 to 12 19.31 2 1 16:02 6 0 0.0000 2 9 l6to 14 19.31 2 1 9:45 7 0 0.0000 2 10 16to16 13.11 3 1 16:34 10 4 0.1250 1 13.21 3 1 12:19 1 0 0.0000 1 13.21 3 1 16:30 5 3 0.0938 1 treatment 10 to 12 means: depleted at 10 a.m., 1 13.21 3 1 10:20 7 2 0.0625 2 13.31 3 1 12:10 1 0 0.0000 nectar measured at 12a.m. 13.31 3 1 16:22 5 1 0.0313 2 13.31 3 1 10:13 7 0.5 0.0156 2 18.11 3 1 12:02 1 7.5 0.2344 . 32 1 18.11 3 1 16:15 5 0 0.0000 1 no value means not observed 18.11 3 1 10:05 7 5 0.1563 1 1 18.123 1 13:23 2 0 0.0000 18.123 1 16:45 6 0 0.0000 1 18.123 1 11:02 7 2 0.0625 1 19.31 3 1 16:03 10 4 0.1250 2 1 13.11 4 1 12:25 1 0 0.0000 1 13.11 4 1 16:35 5 0 0.0000 13.11 4 1 10:32 7 0 0.0000 1 13.21 4 1 16:31 10 0 0.0000 1 13.31 4 1 16:23 10 0 0.0000 1 18.11 4 1 16:17 10 0 0.0000 1 18.12 4 1 16:46 10 2 0.0625 1 19.31 4 1 11:52 1 0 0.0000 2 19.31 4 1 16:04 5 3.5 0.1094 2 19.31 4 1 9:49 7 0 0.0000 2 1 13.11 5 1 16:36 10 0 0.0000 13.21 5 1 12:20 1 0 0.0000 1 13.21 5 1 10:23 7 0 0.0000 1 18.125 1 13:29 2 0 0.0000 1 18.125 1 16:48 6 2 0.0625 1 18.125 1 11:04 7 3.5 0.1094 41 1 18.126 1 16:49 10 1 0.0313 1 18.127 1 13:41 2 0 0.0000 1 18.12 7 1 16:51 6 0 0.0000 1 18.12 7 1 11:10 7 3 0.0938 1 18.12 8 1 16:52 9 0 0.0000 1 18.12 9 1 13:42 2 0 0.0000 1 18.12 9 1 16:52 6 0 0.0000 1 18.129 1 11:17 8 0 0.0000 1 18.12 10 1 16:53 10 0 0.0000 1 18.12 11 1 16:53 6 1 0.0313 1 18.12 11 1 13:43 9 0 0.0000 1 13.11 1 2 12:22 1 0 0.0000 1 13.11 1 2 16:32 5 0 0.0000 1 13.11 1 2 10:28 7 0 0.0000 1 13.21 1 2 16:25 5 9 0.2813 1 13.21 1 2 12:16 8 1.5 0.0469 1 13.31 1 2 16:19 5 0 0.0000 2 13.31 1 2 12:09 8 0 0.0000 2 13.32 1 2 16:38 5 1 0.0313 2 2 13.32 1 2 12:30 8 5.5 0.1719 1 18.11 1 2 16:13 5 0 0.0000 1 18.11 1 2 11:59 8 9 0.2813., 46 2 19.32 1 2 11:54 1 0 0.0000 2 19.32 1 2 16:06 5 0 0.0000 2 19.32 1 2 9:50 7 1 0.0313 13.11 2 2 16:33 10 0 0.0000 1 13.21 2 2 12:17 1 0 0.0000 1 13.21 2 2 16:29 5 3 0.0938 1 13.21 2 2 10:19 7 1.5 0.0469 1 13.31 2 2 12:10 1 0 0.0000 2 13.31 2 2 16:21 5 0 0.0000 2 13.31 2 2 10:12 7 0 0.0000 2 13.32 2 2 12:30 1 0 0.0000 2 13.322 2 16:38 5 0 0.0000 2 13.322 2 10:54 7 0 0.0000 2 18.11 2 2 12:01 1 0 0.0000 1 18.11 2 2 16:14 5 0 0.0000 1 18.11 2 2 10:04 7 0.5 0.0156 1 18.122 2 12:35 2 4 0.1250 1 18.122 2 16:40 6 2 0.0625 1 18.122 2 10:58 7 4 0.1250 1 19.31 2 2 16:02 5 0.5 0.01 56 2 19.31 2 2 11:48 8 0 0.0000 2 13.11 3 2 12:24 1 0 0.0000 1 13.11 3 2 16:34 5 0 0.0000 1 13.11 3 2 10:29 7 7 0.2188 0 1 13.21 3 2 16:30 5 2 0.0625 1 13.213 2 12:19 8 0 0.0000 1 13.313 2 16:22 10 0 0.0000 2 18.113 2 16:15 5 2 0.0625 1 18.113 2 1203 8 3.5 0.1094 32 1 18.123 2 16:45 6 1 0.0313 1 18.123 2 1323 9 2 0.0625 1 19.31 3 2 11:49 2 9 0.2813 2 19.31 3 2 16:03 6 0.5 0.0156 2 19.31 3 2 9:47 7 0 0.0000 2 13.11 4 2 16:35 5 0 0.0000 1 13.114 2 12:25 8 0 0.0000 1 13.21 4 2 12:20 1 0 0.0000 1 13.214 2 16:31 5 0 0.0000 1 13.21 4 2 10:22 7 0 0.0000 1 13.314 2 12:12 1 0 0.0000 1 13.314 2 16:23 5 0 0.0000 1 13.31 4 2 10:14 7 0 0.0000 1 18.11 4 2 12:04 1 0 0.0000 1 18.11 4 2 16:17 5 5 0.1563 1 18.11 4 2 10:06 7 3 0.0938 1 18.124 2 13:24 2 4 0.1250 50 1 18.124 2 16:46 6 0 0.0000 1 18.124 2 11:03 7 3.5 0.1094 1 19.31 4 2 16:04 5 1 0.0313 2 19.31 4 2 11:52 8 0 0.0000 2 1 13.11 5 2 12:28 1 1 0.0313 13.11 5 2 16:36 5 0 0.0000 1 13.11 5 2 10:34 7 0 0.0000 1 13.21 5 2 12:20 8 0 0.0000 1 18.125 2 16:48 6 0 0.0000 1 18.125 2 13:29 9 2 0.0625 1 18.126 2 13:30 2 5 0.1563 1 18.126 2 16:49 6 0 0.0000 1 18.126 2 11:09 7 5 0.1563 1 18.127 2 16:51 6 0 0.0000 1 18.127 2 13:41 9 1 0.0313 1 18.128 2 11:13 6 8 0.2500 46 1 18.128 2 16:52 6 0 0.0000 1 18.128 2 13:41 9 2.5 0.0781 1 18.129 2 16:52 6 1 0.0313 1 18.129 2 13:42 9 2.5 0.0781 1 18.12 10 2 11:17 2 5.5 0.1719 1 18.12 10 2 13:43 4 3 0.0938 1 18.12 10 2 16:53 6 0 0.0000 1 1 13.21 1 3 13:59 2 0 0.0000 1 13.21 1 3 16:25 6 5 0.1563 1 13.21 1 3 10:17 7 0 0.0000 1 13.31 1 3 13:57 2 0 0.0000 2 13.31 1 3 16:19 6 0 0.0000 2 13.31 1 3 10:10 7 1 0.0313 2 13.32 1 3 10:53 7 2.5 0.0781 1 18.11 1 3 13:52 2 0 0.0000 1 18.11 1 3 16:13 6 0 0.0000 1 18.11 1 3 10:03 7 0 0.0000 2 19.32 1 3 16:06 5 0 0.0000 2 19.32 1 3 11:54 8 0 0.0000 13.21 2 3 16:29 5 1 0.0313 1 13.21 2 3 12:17 8 0.5 0.0156 1 18.11 2 3 16:14 5 0 0.0000 1 18.11 2 3 12:01 8 0 0.0000 1 18.12 2 3 16:40 5 3 0.0938 1 18.12 2 3 12:39 8 0.5 0.0156 1 19.31 2 3 13:48 2 0 0.0000 2 19.31 2 3 9:46 6 0 0.0000 2 19.31 2 3 16:02 6 0 0.0000 2 13.11 3 3 16:34 5 0 0.0000 1 13.113 3 12:24 8 0.5 0.0156 1 13.213 3 14:00 2 0 0.0000 1 13.213 3 16:30 6 1 0.0313 1 13.21 3 3 10:20 7 0 0.0000 1 18.11 3 3 13:53 2 1 0.0313turn 1 18.11 3 3 16:15 6 1 0.0313 1 18.11 3 3 10:06 7 0 0.0000 1 18.123 3 16:45 6 1 0.0313 1 18.123 3 14:13 9 0 0.0000 1 19.31 3 3 11:50 8 1.5 0.0469 30 2 13.11 4 3 14:05 2 1 0.0313 1 13.11 4 3 16:35 6 0 0.0000 1 13.11 4 3 10:33 7 0 0.0000 1 13.21 4 3 16:31 5 0 0.0000 1 13.21 4 3 12:20 8 0 0.0000 1 13.31 4 3 16.23 5 0 0.0000 1 13.31 4 3 12:12 8 0 0.0000 1 18.114 3 16:17 5 3 0.0938 1 18.11 4 3 12:04 8 3 0.0938 1 18.124 3 16:46 6 1 0.0313 1 18.124 3 13.24 9 4 0.1250 50 1 13.11 5 3 16:36 5 0 0.0000 1 13.11 5 3 12:28 8 1.5 0.0469 1 13.21 5 3 14:01 2 1.5 0.0469 1 13.21 5 3 10:23 7 3 0.0938 1 18.125 3 14:15 2 2 0.0625 1 18.125 3 16:48 6 1 0.0313 1 18.12 5 3 11:05 7 1 0.0313 41 1 18.126 3 16:49 6 1 0.0313 1 18.126 3 13:31 9 1.5 0.0469 1 18.127 3 14:16 2 3.5 0.1094 1 18.127 3 16:51 6 1 0.0313 1 18.127 3 11:11 7 0.5 0.0156 1 18.128 3 16:52 6 0 0.0000 1 18.128 3 13:42 9 0.5 0.01 56 1 18.129 3 14:19 2 0 0.0000 1 18.129 3 16:52 6 0 0.0000 1 18.129 3 11:17 8 2.5 0.0781 1 18.12 10 3 16:53 6 0 0.0000 1 18.12 10 3 13:43 9 0 0.0000 1 1 13.21 1 4 16:25 6 2 0.0625 1 13.21 1 4 13:59 9 0 0.0000 2 13.31 1 4 16:19 6 0 0.0000 1 13.31 1 4 13:57 9 0 0.0000 1 18.11 1 4 16:13 6 1 0.0313 1 18.11 1 4 13:52 9 2 0.0625 2 19.32 1 4 13:49 2 0 0.0000 2 19.32 1 4 16:06 6 0 0.0000 2 19.32 1 4 9:51 7 6.5 0.2031 13.21 2 4 16:29 3 1 0.0313 1 13.21 2 4 10:19 7 2 0.0625 1 18.122 4 14:10 2 0 0.0000 1 18.122 4 16:40 6 1 0.0313 1 18.122 4 10:59 7 1.5 0.0469 1 13.113 4 16:34 3 1 0.0313 1 13.113 4 10:30 7 0.5 0.0156 0 1 13.21 3 4 16:30 6 0 0.0000 1 13.21 3 4 14:00 9 1 0.0313 1 18.11 3 4 13:53 9 2.5 0.0781 turn 1 13.11 4 4 16:35 6 0 0.0000 1 13.11 4 4 1405 9 0.5 0.0156 1 13.21 4 4 16:31 3 0 0.0000 1 13.21 4 4 10:22 7 6.5 0.2031 1 13.31 4 4 16:23 3 3 0.0938 1 13.31 4 4 10:15 7 0 0.0000 1 18.11 4 4 16:17 3 3 0.0938 1 18.11 4 4 10:08 7 0 0.0000 1 18.12 4 4 14:14 2 3 0.0938 1 18.12 4 4 16:46 6 0 0.0000 1 18.12 4 4 11:04 7 0.5 0.0156 1 13.11 5 4 16:36 3 0 0.0000 1 13.11 5 4 10:34 7 0.5 0.0156 1 13.21 5 4 14:01 9 0 0.0000 1 18.125 4 16:48 6 0 0.0000 1 18.125 4 14:15 9 0.5 0.0156 1 18.127 4 16:51 6 0 0.0000 1 18.127 4 14:17 9 1 0.0313 1 18.128 4 14:18 2 1.5 0.0469 1 18.128 4 16:52 6 0 0.0000 1 18.128 4 11:17 8 2.5 0.0781 46 I 1 18.129 4 16:52 10 0 0.0000 1 13.31 1 5 12:09 1 0 0.0000 2 13.31 1 5 16:19 5 2 0.0625 2 13.31 1 5 10:11 7 1 0.0313 2 13.21 2 5 16:29 10 0 0.0000 1 18.122 5 16:41 6 0 0.0000 1 18.122 5 14:10 9 0 0.0000 1 13.11 3 5 16:34 6 0 0.0000 1 13.11 3 5 14:03 9 1.5 0.0469 1 13.21 3 5 16:30 5 1 0.0313 1 1321 3 5 12:19 8 0 0.0000 1 13.11 4 5 12:25 1 0 0.0000 1 13.11 4 5 14:05 4 0.5 0.0156 1 13.11 4 5 16:35 6 0 0.0000 1 13.11 4 5 10:33 7 1 0.0313 1 13.21 4 5 14:01 9 2 0.0625 1 13.31 4 5 16:23 10 3 0.0938 1 18.12 4 5 16:46 6 0 0.0000 1 18.12 4 5 14:14 9 1 0.0313 1 18.12 5 5 13:29 2 0 0.0000 1 18.125 5 16:48 6 0 0.0000 1 18.125 5 11:05 7 1 0.0313 41 1 18.127 5 13:41 2 0 0.0000 1 18.127 5 16:51 6 0 0.0000 1 18.127 5 11:11 7 1 0.0313 1 18.128 5 16:52 6 1 0.0313 1 18.128 5 14:19 9 0.5 0.0156 1 18.129 5 16:52 10 0 0.0000 1 13.31 1 6 16:19 5 1 0.0313 2 13.31 1 6 12:09 8 0 0.0000 2 13.21 2 6 16:29 10 0 0.0000 1 18.122 6 12:41 1 0 0.0000 1 18.122 6 16:41 5 0 0.0000 1 18.122 6 11:00 7 0.5 0.0156 1 13.213 6 12:19 8 0 0.0000 1 13.114 6 16:35 5 2 0.0625 1 13.11 4 6 12:25 8 1.5 0.0469 1 18.12 4 6 16:46 6 0 0.0000 1 18.12 4 6 13:24 9 2 0.0625 50 1 18.12 5 6 16:48 6 0 0.0000 1 18.125 6 13:30 9 5 0.1563 1 18.12 7 6 16:51 6 0 0.0000 1 18.12 7 6 13:41 9 2 0.0625 1 13.31 1 7 16:19 6 0 0.0000 2 13.31 1 7 13:57 9 2 0.0625 1 18.122 7 16:41 5 1 0.0313 1 18.122 7 12:41 8 0 0.0000 1 18.125 7 14:16 2 0.5 0.0156 1 18.125 7 16:48 6 0 0.0000 1 18.125 7 11:06 7 10.5 0.3281 41 1 18.127 7 14:18 2 0 0.0000 1 18.127 7 16:51 6 0 0.0000 1 18.127 7 11:12 7 0.5 0.0156 1 13.31 1 8 16:19 10 0 0.0000 2 18.12 2 8 14:11 2 1 0.0313 1 18.12 2 8 16:41 6 0 0.0000 1 18.122 8 11:01 7 0.5 0.0156 1 18.125 8 16:48 6 0 0.0000 1 18.125 8 14:16 9 0.5 0.0156 1 18.127 8 16:51 6 0 0.0000 1 18.127 8 14:18 9 0 0.0000 1 13.31 1 9 16:19 10 1 0.0313 2 18.122 9 16:42 6 1 0.0313 1 18.122 9 14:11 9 2.5 0.0781 1 18.127 9 13:41 2 1 0.0313 1 18.127 9 16:51 6 0 0.0000 1 18.127 9 11:12 7 0.5 0.0156 1 18.122 10 12:41 1 2.5 0.0781 1 18.12 2 10 16:42 5 1 0.0313 1 18.122 10 1t02 7 0 0.0000 1 18.122 11 16:42 5 1 0.0313 1 18.122 11 12:41 8 0 0.0000 1 18.12 2 12 14:12 2 0.5 0.0156 1 18.122 12 16:42 6 0 0.0000 1 18.122 12 11:02 7 0 0.0000 1 date treat- potwhorlflower mm. microliter sugar wvmmcji ment nectar t conc. AppendixI 2001051b iutol2 1 1 1 3 O.lbIb 2001051610 to 12 2 1 1 15 0.9375 nectar Lamium album 20010516lOtol2 3 1 1 3 0.1875 20010516 lOtol2 4 1 1 14 0.8750 51 plants: Manjas 20010516 10 to 12 3 1 6 0 0.0000 source of 40 2001051610to 12 1 2 1 16 1.0000 plants hadbeen netted intheexperimental garden 20010516 lOtol2 2 2 1 19 1.1875 28 20010516 10 to 12 3 2 1 15 0.9375 from 1505200116:00 plants in greenhouse 20010516 lOtol2 4 2 1 16 1.0000 44 20010516 lOto 12 5 2 1 3 0.1875 date of measurements: 1605 2001 20010516 lOtol24 2 6 11 0.6875 42 4 0.2500 2001051610 to 12 5 3 1 * in 2 microliter capillary, 32 mm 20010516lOtol2 5 2 6 5 0.3125 4 2001051610 to 14 1 1 3 0.2500 conc.= concentration 20010516 lOtol4 2 1 3 28 1.7500 41 20010516 lOtol4 3 1 3 47 2.9375 36 20010516 lOtol4 4 1 3 9 0.5625 20010516 lOtol4 5 1 4 8 0.5000 20010516 lOtol4 3 1 8 3 0.1875 37 20010516 lOtol4 1 2 3 47 2.9375 Treatment legend 2001051610 to 14 2 2 3 6 0.3750 20010516 lOtol4 3 2 3 18 1.1250 41 1 lOtol2 20010516 lOtol4 4 2 3 17 1.0625 47 2 101014 20010516 lOtol4 5 2 3 0 0.0000 3 lOtol6 20010516 lOtol4 4 2 8 48 3.0000 41 4 12to14 20010516 101014 5 2 8 2 0.1250 5 121016 20010516 lOtol4 5 3 3 31 1.9375 39 6 7 l6tolO 20010516 lOtol6 5 1 1 5 0.3125 8 16to12 20010516 lOtol6 1 1 5 94 5.8750 32 9 161014 20010516 lOtol6 2 1 5 52 3.2500 31 10 161016 20010516 1010163 1 5 15 0.9375 30 20010516 1010164 1 5 20 1.2500 42 20010516 lOtol6 5 1 6 2 0.1250 treatmentlotol2means:depletedatloa.m. 20010516 101016 3 1 10 24 1.5000 33 20010516 lOtol6 1 2 5 45 2.8125 21 nectarmeasuredat 12a.m. 20010516 1010162 2 5 3 0.1875 20010516 lOtol6 3 2 5 14 0.8750 26 20010516 lOtol64 2 5 22 1.3750 37 20010516 lOtol6 5 2 5 18 1.1250 37 no value means not observed 20010516 101016 4 2 10 29 1.8125 32 I 20010516 12to14 1 1 1 0 0.0000 20010516 12to14 2 1 1 12 0.7500 20010516 12to14 3 1 1 2 0.1250 20010516 12to144 1 1 26 1.6250 50 20010516 12to14 1 2 1 41 2.5625 36 20010516 12to14 2 2 1 11 0.6875 20010516 12to143 2 1 1 0.0625 20010516 12to14 4 2 1 25 1.5625 40 20010516 12to14 5 2 1 1 0.0625 20010516 12to14 4 2 6 25 1.5625 43 20010516 12to14 5 2 6 4 0.2500 20010516 12to14 5 3 1 2 0.1250 20010516 12to16 1 1 2 11 0.6875 20010516 121016 3 1 2 3 0.1875 20010516 12to16 4 1 2 28 1.7500 45 20010516 12to16 5 1 2 20 1.2500 40 20010516 12to16 3 1 7 31 1.9375 35 20010516 12to16 1 2 2 15 0.9375 14 20010516 12to16 2 2 2 20 1.2500 21 20010516 12to16 3 2 2 3 0.1875 20010516 12to16 4 2 2 8 0.5000 20010516 12to16 5 2 2 12 0.7500 20010516 12to16 4 2 7 25 1.5625 29 20010516 12to16 2 1 2 18 1.1250 15 20010516 14to16 1 1 1 0 0.0000 20010516 14to16 2 1 1 2 0.1250 20010516 141016 3 1 1 3 0.1875 20010516 14to164 1 1 9 0.5625 20010516 141016 1 1 4 43 2.6875 45 20010516 14to16 2 1 4 8 0.5000 20010516 14to16 4 1 4 3 0.1875 20010516 14to16 5 1 4 7 0.4375 20010516 14to16 3 1 9 3 0.1875 20010516 14to16 1 2 1 2 0.1250 20010516 14to16 2 2 1 3 0.1875 20010516 141016 3 2 1 0 0.0000 20010516 14to16 4 2 1 3 0.1875 20010516 14to16 5 2 1 0 0.0000 0.0000 20010516 14to16 1 2 4 0 20010516 14to16 2 2 4 12 0.7500 20010516 14to16 3 2 4 2 0.1250 20010516 14to16 4 2 4 31 1.9375 40 20010516 14to16 5 2 4 47 2.9375 51 20010516 14to16 4 2 6 15 0.9375 32 20010516 141016 4 2 9 20 1.2500 40 1 4 0.2500 20010516 l6tolO 1 1 0.1250 20010516 l6tolO 3 1 1 2 0.5313 20010516 l6tolO 4 1 1 8.5 0.3125 20010516 l6tolO 5 1 1 5 1.5625 50 20010516 l6tolO 1 1 3 25 0.0000 20010516 l6tolO 2 1 3 0 0.9375 44 20010516 l6tolO 3 1 3 15 0.7500 20010516 l6tolO 4 1 3 12 20010516 l6tolO 5 1 3 9 0.5625 0.3438 20010516 l6tolO 1 1 5 5.5 1.5000 41 20010516 l6tolO 2 1 5 24 15 0.9375 49 20010516 l6tolO 3 1 5 0.9375 50 20010516 l6tolO 4 1 5 15 0.0000 20010516 l6tolO 5 1 5 0 0.1563 20010516 l6tolO 3 1 6 2.5 1 0.0625 20010516 l6tolO 3 1 8 17 1.0625 51 20010516 l6tolO 3 1 10 1.8750 44 20010516 l6tolO 1 2 1 30 0.0000 20010516 l6tolO 2 2 1 0 1.3750 39 20010516 l6tolO 3 2 1 22 1.0000 20010516 l6tolO 4 2 1 16 3.3125 48 20010516 l6tolO 1 2 3 53 20010516 l6tolO 2 2 3 56 3.5000 35 20010516 l6tolO 3 2 3 12 0.7500 20010516 l6tolO 4 2 3 13 0.8125 20010516 l6tolO 5 2 3 0 0.0000 52 3.2500 37 20010516 l6tolO 1 2 5 20010516 l6tolO 2 2 5 22 1.3750 24 20010516 l6tolO 3 2 5 18 1.1250 32 20010516 l6tolO 4 2 5 17 1.0625 42 20010516 l6tolO 5 2 5 8 0.5000 20010516 l6tolO 4 2 6 16 1.0000 47 5 2 6 14 0.8750 20010516 l6tolO 44 20010516 l6tolO 4 2 8 22 1.3750 20010516 l6tolO 5 2 8 12 0.7500 20010516 l6tolO 4 2 10 0 0.0000 20010516 l6tolO 5 2 10 14 0.8750 20010516 l6tolO 5 3 1 9 0.5625 20010516 l6tolO 5 3 3 15 0.9375 20010516 l6tolO 5 3 5 0 0.0000 3.3125 45 20010516 l6tolO 2 1 1 53 18 1.1250 52 20010516 16to12 1 1 2 12 0.7500 20010516 16to12 2 1 2 10 0.6250 20010516 16to123 1 2 1.1875 52 20010516 161012 4 1 2 19 1.5625 50 20010516 16to12 5 1 2 25 32 2.0000 50 20010516 161012 3 1 7 13 0.8125 20010516 161012 1 2 2 20010516 161012 2 2 2 14 0.8750 29 20010516 16to12 3 2 2 18 1.1250 31 20010516 16to124 2 2 10 0.6250 20010516 16to12 5 2 2 21 1.3125 52 20010516 16to12 4 2 7 17 1.0625 44 47 20010516 16to12 5 2 7 21 1.3125 20010516 16to12 5 3 2 25 1.5625 50 19 1.1875 50 20010516 16to14 1 1 4 1.2500 35 20010516 16to14 2 1 4 20 0.1875 20010516 16to14 3 1 4 3 3.1250 51 20010516 16to14 4 1 4 50 1 0.0625 20010516 16to143 1 9 18 1.1250 40 20010516 16to14 1 2 4 34 20010516 16to14 2 2 4 18 1.1250 43 20010516 16to14 3 2 4 18 1.1250 48 20010516 16to14 4 2 4 32 2.0000 52 20010516 161014 5 2 4 16 1.0000 20010516 16to14 4 2 9 27 1.6875 51 52 20010516 16to14 5 3 4 18 1.1250 52 20010516 16to14 5 2 9 31 1.9375 0.2500 20010516 161016 5 1 7 4 1.9375 45 20010516 161016 5 1 9 31 0.3125 20010516 16to16 5 1 10 5 20010516 161010 5 2 1 0 0.0000 ______# # # flowers % visited % visited % visited total avg. # total yyyymmdd date verge number plot insect species visitedtypeplant (sec)staytime 77 visited visitedintl. 3 fi.4 on visited infl. available 6 f I. of fl. availableon visited intl. 66.7 # intl. infl.In plot of total 2.19 # f I. infI. plot of total 1.16 in plot# inf. 137 in plottl/inf.2.52 in plot 345# fi. 200105082001050320010509 19.5 18 5 683 ApisApidae mellifera small hermaphroditicfemale 1803283 16 35 3110 5 4215 9 55.673.866.7 29.090.619.09 16.010.325.17 49055 3.163.52 1548194 2001050920010508 19.5 18 768 Apis mellifera femalehermaphroditic 194751610 30 624 41 62 1031012 5 60.040.050.039.8 0.406.121.202.92 0.510.172.651.74 500490137 2.362.523.16 11801548345 20010509 18 7 Apis mellifera femalefemale 702084 18 96 2612 8 532319 42.149.152.2 3.601.801.20 2.200.681.02 500 2.36 11801180 20010509 18 7 Apis mellifera female 154130120 179 6 11 54.5 3.401.80 0.51 500500 2.362.36 1180 2001050320010509 19.540 13188 BombuspescuorumApis me/hf era femalehermaphroditic 3319 5 34 1 471 13 3 53.833.330.8 3.450.560.82 0.160.261.92 18049087 4.203.523.16 1548365634 20010510 27 2 13 1 Bombuspascuorum female 96 14 1 10.371.15 13587 2.484.20 335365 200105092001051020010509 2 1 Bombuspascuorum hermaphroditic 11227 24 9 33 60 55.0 17.786.67 9.86 135 2.48 335 20010510 3417 15 15 HymenopteraBombuspascuorum (koekoekshommel) female 10 7 2 1 6 7 85.7 2.411.54 3.69 6583 3.561.96 231163 20010503 2 1 Melanostoma sp./Platycheirussp./Platycheinis sp. hermaphroditic 7068 3 1 1 2 50.0 0.742.22 0.30 135 2.48 335 200105032001 0503 22 1 Melanostoma sp.,r'IatycheirusspiPlatycheirus sp. hermaphroditic 14483 3 1 3 12 94 50.033.3 0.742.22 0.600.90 135 2.48 335335 2001050320010508 19.5 17 2 85 1 Melanostoma sp./Platycheirussp.IPlatycheirusspiPlatycheirus sp. hermaphroditicfemale 118373 2 31 42 12 38 33.366.712.5 0.610.741.20 0.260.301.23 49013583 3.162.481.96 1548163335 20010508 25 10 Melanostoma sp.iPlatycheirusno value sp. means: not observed female 68 1 3 5 differentfi. AppendixPlot= flowers; observations plots mt. were = inflorescences J given(10 mm) a number: on Glechoma plot number hederacea60.0 1.37 1.27 73 3.24 237 1 15 2001050920010509 22 1 MelanostomaM&anostomasp./PlatyCheirUSSP. sp./Platycheirus sp. hermaphroditic 16 1 1 1 42 50.025.0 0.740.74 0.30 135 2.48 335 20010509 17 2 4 Melanostoma sp./Platycheirus sp. femalehermaphroditic 26 4 1 1 4 25.0 0.91 0.52 110110 1.761.76 194 2001050920010509 17 54 1 MelanostornaMelanostoma sp.IPlatycheirussp./Platycheirus sp. female 4165 1 1 4 25.0 1.20 0.61 83 1.96 261163 20010510 2 1 Melanostoma sp./P!atycheirussp./Platycheinis sp. female 154 1 1 25.033.3 1.16 0.38 86 3.04 261 2001050320010510 22 1 1 MelanostomaRhingia campestris sp./Platycheirus sp. femalehermaphroditic 32 2 2 5 40.0 1.481.16 0.600.76 13586 2.483.04 335261 20010503 26.5 17 125 Rhingia campestris hermaphroditicfemale 6716 2 1 2 1 84 50.012.5 0.972.41 0.261.23 10383 3.681.96 379163 20010508 2517 104 Rhingia campestnscampestris female 2025 1 1 100.050.0 0.911.37 0.420.52 11073 3.241.76 237194 20010508 26 11 RhingiacampestrlsRhingia campestris hermaphroditic 4629 5 1 5 1 17 7 28.629.4 0.974.85 0.531.32 103 3.68 379 2001050920010508 26.5 17 12 4 RhingiaRhingia campestns campestns femalehermaphroditic 51 96 2 1 53 40.033.3 1.820.97 0.261.03 110103 3.681.76 379194 20010510 17 54 RhlngiaRhingia campestriscampestns female 1 1 2 12 50.0 0.911.20 0.610.52 83 1.961.96 163 20010510 19.5 17 58 RhingiaRhingiacampeStl7S campestns hermaphroditicfemale 2715 4 1 1 84 100.025.012.5 0.202.27 0.450.06 490 44 5.083.16 1548224 2001051020010510 27.527.5 1614 RhingiaRhingia campestris campestris femalehermaphroditicfemale 1220 1 1 34 25.066.7 2.862.27 2.420.45 3544 2.365.08 22483 2001050320010509 34 25 2 1 SyrphidaeSyiphidae blackyellow hermaphroditic 5 1 4 25.0 0.74 0.30 13585 1.882.48 335160iso 2001050320010508 19.5 40 5 18329 femalefemale 47518055 4.963.52i. 2356634194 20010508 25 10 female 73 3.24 237 20010509 19.5 342 169 1 hermaphroditIchermaphroditicfemale 4753586 4.002.364.963.04 2356261 83 200105092001051020010509 27.5 3440 151417 femalefemalehermaphroditic 4465 3.565.08 231224 yyyymmdd date verge time hightypeplot speciesinsect stay time (sec) 37 # visitedumbels 2 # visitedumb.es 3 # availableon visited umb.es umbels 18 total # umbels in 19 plot umb.es/umbel avg. 8.64 164.09in plot 20010605 3535 11:0010:50 low EmpisMuscaEmpis tessellatasp. tessellata 600600 4 1 10 40 23 8.69 199.92 20010605 35 11:00 low Empis tessellata 235329 23 1 23 8.698.69 199.92 20010605 35 11:00 low Eristalissp.Empis tessellata 15339 4 1 5 38 2323 8.69 199.92 20010605 35 11:00 low MuscaEristalls tenax/perlinax sp. 32120 3 1 3 9 23 8.69 199.92 20010605 35 11:1011:00 highlow MuscaEmpis sp. tessellata 536140 5 1 29 2 4310 192319 8.64 164.09 20010605 35 11:10 high TachinaEmpis tessellata fabricia 14593 13 1017 2410 20 8.698.64 173.85164.09 20010605 35 11:20 low EristalishoilicolaEristalisEmpistessollata horticola 52015040 2 1 51 1810 20 8.69 173.85 20010605 35 11:20 low Eristalis tenax/pertinax 4586 8 1 23 1 10 20 8.69 173.85 20010605 35 11:20 low MuscaRhingiaMuscasp. sp. campestris 121 7 12 4 1 1017 20 8.69 173.85 20010605 3535 11:20 low Sarcophaga sp. 14057 4 1 11 2 1039 8.69 173.85 20010605 35 12:00 low2 CanthansCanthanssp. sp. 139155 1 83 9 20 10.40 208.00 20010605 35 12:00 low2 Empis diagramma 199155 1 2 10 20 10.4010.40 208.00208.00 20010605 35 12:00 low2 Empis dlagrammadiagramma 166136101 21 24 3 10 20 10.40 208.00 20010605 35 12:1012:00 high2hlgh2low2 EristalisarbustorumEmplstessellata Musca sp. 28517821 52 1 2419 3 472211 20 11.83 236.67 AppendixPlot observations K(10 mm) 20010605 35 12:1012:20 high2low2 EnstalisEristalishortlcola Empis arbustorum diagramma 152 1 2 10 9 20 11.8310.40 208.00236.67 plotumb.esAnthnscus type = umbellules sylvestris = location in 2001060520010605 35 12:3012:20 hlgh2low2 DipteraEmpistessellata Empis small diagramma 16312444 21 1015 8 1110 20 11.8310.40 236.67208.00 no values = not observed Appendix L no value meansnot(hlng) observed nsect observations Glechoma hederacea inf.=inflorescences
date verge time plant species 41 insectspecies vvwmrndd nt. insects 2uui uu 2 15:30 Cardaminepraterisis 12 Metanostoma sp.JPlatycheirus sp. 20010502 2 15:30 Glechoma hederacea female 110 Bombus pascuorum 20010502 2 15:30 Glechomahederacea female 110 machis 10 1 20010502 2 15:30 Glechomahederaceahermaphroditic 600 Bombuspascuorum 20010502 2 15:30 Taraxacum officinale 12 Sy,phidae 20010502 5 15:00-15:20 Syrphidae(hommelzw) 20010502 5 15:00-15:20 Hymenoptera(koekhom) 2 20010502 5 15:00-15:20 Taraxacum officinale 89 Formicoidea 20010502 5 15:00-15:20 Taraxacum officinale 89 Apis mellifera 20010502 5 15:00-15:20 Taraxacum officinale 89 Apidaesmall 20010502 5 15:00-15:20 Taraxacum officinale 89 Enstalis arbustorum 1 20010502 9 14:45 Glechomahederacea female 5 Bombus pascuorum 1 20010502 9 14:45 Taraxacumofficinale 75 Apismellifera 1 20010502 9 14:45 Taraxacum officinale 75 Eristalis sp. 2 20010502 9 14:45 Taraxacumofficinale 75 Helophilus sp. 1 20010502 9 14:45 Taraxacum officinale 75 Scatophaçjastercoraria 20010502 12 14:00 Taraxacumofficinale 194 Formicoidea 20010502 12 14:00 Taraxacum off icinale 194 Apismellifera 1 20010502 12 14:00 Taraxacumofficinale 194 Bibiosp. 4 20010502 12 14:00 Taraxacum officinale 194 Enstalissp. 1 20010502 17 13:45-13:50 Glechomahederacea hermaphroditic 348 Bombus pascuorum 20010502 17 13:45-13:50 Taraxacum officinale 367 Bibio sp. 6 20010502 17 13:45-13:50 Taraxacum off icinale 367 Gymnochaeta vindis 20010502 17 13:45-13:50 Taraxacum officinale 367 Scatophapastercorana 24 20010502 17 13:45-13:50 Taraxacum officinale 367 Syrphidae 20010502 18 13:30-13:35 GleeMoma hederacea female 1552 Apis mellifera 3 20010502 25 11:00 Taraxacum officinale 78 Formicoidea 3 20010502 25 11:00 Taraxacum officinale 78 Biblo sp. 2 20010502 26 11:30 Taraxacum officinale 108 Apis mellifera 1 20010502 26 11:30 Taraxacum officinale 108 Bibio sp. 1 20010503 19.5 14:15 Glechomahederacea female 400 Apis mellifera 4 20010503 19.5 14:15 Glechomahederaceafemale 400 Melanostoma spiPlatycheirus sp. 2 20010503 19.5 14:15 Glechoma hederacea hermaphroditic 5000 Apis mellifera 1 20010503 19.5 14:15 Glechomahaderaceahermaphroditic 5000 Bombus pascuorum 4 20010503 19.5 14:15 Glechoma haderacea hermaphroditic 5000 Eristalissp. 20010503 19.5 14:15 Glechoma hede,aceahermaphroditic 5000 Helophilus sp. 20010503 19.5 14:15 Glechomahederaceahermaphroditic 5000 Melanostoma sp./Platycheirus sp. 20010503 19.5 14:15 Glechomahederacea hermaphroditic 5000 Rhinqia campestris 7 20010508 17 10:45-11:08 Glechomahederacea hermaphroditic 473 Bombus pascuorum 20010508 17 10:45-11:08 Glechoma hederaceahermaphroditic 473 Bombus pascuorum 20010508 17 10:45-11:08 Glechoma hederacea hermaphroditic 473 Melanostomasp./Platycheirus sp. 20010508 17 10:45-11:08 Glechoma hederacea hermaphroditic 473 Rhingia campestris 2 20010508 17 10:45-11:08 Glechomahederaceahermaphroditic 473 Dipterasmall brown 2 20010508 17 10:45-11:08 Glechomahederacaa hermaphroditic 473 Diptera small black 2 20010508 18 14:01 Taraxacum officinale 1598 Diptera small I 20010508 18 14:01 Taraxacumofficinale 1598 Diptera small 1 20010508 18 14:01 Taraxacumofficinale 1598 Dipterasmall 8 20010508 18.5 14:01 Glechomahederacea female 1552 Bombuspascuorum 1 20010508 18.5 14:01 Glechoma hederacea female 1916 Bombussp. 1 20010508 18.5 14:01 Lamiumalbum 281 Bombus pascuofum 2 20010508 18.5 14:01 Lamiumalbum 281 Bombus terrestris 1 20010508 18.5 14:01 Lamium album 281 Rhingia campestns 2 20010508 19.515:27-15:35 GlecMoma hederacea hermaphroditic 5840 Bombussp. 1 20010508 19.515:27-15:35 Glechomahederacea hermaphroditic 5840 Helophilus sp. 2 20010508 19.515:27-15:35 Glechoma hederaceahermaphroditic 5840 Rhingia campestns 11 20010509 2 10:24 Glechoma hederaceahermaphroditic 690 Melanostoma sp./Platvcheirus sp. 4 20010509 2 10:24 Glechomahederacea hermaphroditic 690 Melanostoma spiPlatycheirus sp. 20010509 5 15:13 Glechomahederacea hermaphroditic 0 20010509 40 12:11 Glechoma hederaceahermaphroditic 760 Bombus pascuorum 20010510 26 13:57-14:09 Glechoma hederacea female 15 0 20010510 26 13:57-14:09 Glechomahederacea hermaphroditic 65 Rhinqia campestns 20010510 26.513:57-14:09 Glechomahederacea female 4 0 20010510 26.513:57-14:09 Glechomahederacea hermaphroditic 76 0 20010510 27 11:38-11:41 Glechomahederaceafemale 95 0 20010510 27 11:38-11:41 Glechomahederaceahermaphroditic 0 0 20010510 27.511:42-11:44 Glechomahederaceafemale 49 0 20010510 27.511:42-11:44 Glechomahederacea hermaphroditic 0 20010510 34 9:32 Glechomahederaceafemale 154 0 20010510 34 9:32 Glechomahederaceahermaphroditic 42 0 20010510 25+25.f 14:50-15:05 Glechoma hederacea female 159 0 20010510 25+25.f 14:50-15:05 Glechomahederaceahermaphroditic 51 Rhinqia campestris date verqe time # insect species # insects yvvymmdd s=south umbels IAppend1xM n=north I observations Anthriscus s}dvesf 115 20010605 35 9:51 820 Col000teraqreefl A 20010605 35 9:51 820 Elateridae All'erv viSi I s were on .sy yes ns 20010605 35 9:51 820 Elateridae 1 20010605 35 9:51 820 Empis small 2 20010605 35 9:51 820 Empis diaqramma 20010605 35 9:51 820 Empis tessellata 10 no value means:not counted 20010605 35 9:51 820 Eristalis horticola 3 20010605 35 9:51 820 Eristalis tenax/pertinax 7 20010605 35 9:51 820 Musca sp. 3 20010605 35 9:51 820 Scatophaga stercoraria 5 20010605 35 9:51 820 Scatophaqax 1 20010605 35 9:51 820 Symphyta 1 20010605 35 10:03 820 Cantharis so. 5 20010605 35 10:03 820 Cantharis so. 1 20010605 35 10:03 820 Diptera smallblack 1 20010605 35 10:03 820 Elateridae 1 20010605 35 10:03 820 Empis small 3 20010605 35 10:03 820 Empis diapramma 3 20010605 35 10:03 820 Empis tessellafa 7 20010605 35 10:03 820 Eristalis arbuston.im 2 20010605 35 10:03 820 Eristalis tenaxipertinax 2 20010605 35 10:03 820 Maniola jurtina 20010605 35 10:03 820 Musca sp. 20010605 35 10:03 820 Sarcophaqa sp. I 20010605 35 10:03 820 Scatophaqa stertoraria 11 20010605 35 10:03 820 Scatophaqa x 20010605 35 10:03 820 Symphyta larrje 20010605 35 10:03 820 Tachina fabncia 1 20010529 20.5n 11:06 Cantharis sp. 2 20010529 20.5n 11:06 Coleoptera (rozenkever) 1 20010529 20.5n 11:06 Coleoptera oranqe 20010529 20.5n 11:06 Diptera small black 3 20010529 20.5n 11:06 Diptera small brown 3 20010529 20.5n 11:06 Empis tessellata 20010529 20.5n 11:06 Heteroptera 1 20010529 20.5n 11:06 Musca domestica 1 20010529 20.5n 11:06 Musca domestica 3 20010529 20.5n 11:06 Scatophaqa stercoraria 1 20010529 20.5n 11:06 Syrphidae small 1 20010529 20.5n 12:00 Biblo sp. 2 20010529 20.5n 12:00 Cantharis so. 1 20010529 20.5n 12:00 Canthans sp. 4 20010529 20.5n 12:00 Cantharis sp. 2 20010529 20.5n 12:00 Diptera small brown 1 20010529 20.5n 12:00 Diptera v 2 20010529 20.5n 12:00 Elateridae 3 20010529 20.5n 12:00 Empis tessellata 3 20010529 20.5n 12:00 Eristalis tenax/pertinax 1 20010529 20.5n 12:00 Helophilus trivittatus 2 20010529 20.5n 12:00 Tipulldae 1 20010529 20.5n 12:00 Musca sp. 20010529 20.5n 12:00 Symphyta black 2 20010529 20.5n 13:20 Cantharis sp. 20010529 20.5n 13:20 Cant ha ris sp. 1 20010529 20.5n 13:20 Diptera small black 1 20010529 20.5n 13:20 Diptera small brown 2 20010529 20.5n 13:20 Diptera y 1 20010529 20.5n 13:20 Empis tessellata 4 20010529 20.5n 13:20 Eristalis tenaxlpertinax 1 20010529 20.5n 13:20 Heteropt era 1 20010529 20.5n 13:20 Musca sP. 2 20010529 20.5n 13:20 Scatophaqa stercoraria 1 20010529 20.5n 13:20 Symphyta red back 1 20010529 20.5n 14:05 Canthans sp. 4 20010529 20.5n 14:05 Cantharissp. 1 20010529 20.5n 14:05 Elateridae 2 20010529 20.5n 14:05 Empis tessellata 10 20010529 20.5n 14:05 Eristalis arbustorum 2 20010529 20.5n 14:05 Eristalis tenaxlpertinax 5 20010529 20.5n 14:05 Syrphidae (hommelzw) 1 20010529 20.5n 15:05 Araschnia levana prorsa 1 20010529 20.5n 15:05 Cantharis sp. 4 20010529 20.5n 15:05 Empis lMda 1 20010529 20.5n 15:05 Formicoidea 1 20010529 20.5n 15:05 Heteroptera 1
J 20010529 20.5n 15:05 Muscaso. 1 20010529 20.5n 15:05 Muscasp. 2 20010529 20.5n 15:05 Scatophaa stercoraria 4 20010529 20.5n 15:05 Scatophaqax 1 20010529 20.5n 15:05 Curculionidaeqrev 1 20010529 20.5n 15:05 Syrphidae giant 1 20010529 20.5n 15:05 Syrphidae small 1 20010529 20.5$ 11:05 Bibiosp. 1 20010529 20.5s 11:05 Bibio sp. 2 20010529 20.5s 11:05 Canthanssp. 1 20010529 20.5s 11:05 Canthanssp. 2 20010529 20.5s 11:05 Elatenidae 1 20010529 20.5s 11:05 Empis livida 1 20010529 20.5s 11:05 Empis tessellata 5 20010529 20.5s 11:05 Eristalis tenaxlpertinax 4 20010529 20.5s 11:05 Helophilus fri viftafus 1 20010529 20.5$ 11:05 Musca . 4 20010529 20.5s 11:05 Scat ophaçja x 4 20010529 20.5s 11:05 Symphyta black 1 cm 1 20010529 20.5s 11:05 Tachinafabncia 1 20010529 20.5s 12:05 Canthanissp. 1 20010529 20.5s 12:05 Curculionidae 1 20010529 20.5s 12:05 Dipt era small brown 1 20010529 20.5s 12:05 Empis fessellafa 6 20010529 20.5s 12:05 Eristalis horticola 3 20010529 20.5s 12:05 Erisfalis tenax/pertinax 2 20010529 20.5$ 12:05 Heteroptera 1 20010529 20.5s 12:05 Musca sp. 3 20010529 20.5$ 13:20 Canthanis sp. 1 20010529 20.5$ 13:20 Cantharis sp. 2 20010529 20.5$ 13:20 Dipt era small black 2 20010529 20.5s 13:20 Elatenidae 4 20010529 20.5$ 13:20 Empis livida 1 20010529 20.5$ 13:20 Empis tessellata 7 20010529 20.5s 13:20 Eristalisarbustorum 1 20010529 20.5$ 13:20 Eristalishorficola 2 20010529 20.5$ 13:20 Enistalis tenaxlpertinax 14 20010529 20.5s 13:20 Helopiiilus trivittatus 2 20010529 20.5$ 13:20 Muscaso. 6 20010529 20.5s 13:20 Sarcophaqa sp. 2 20010529 20.5$ 13:20 Tachina fabricia 1 20010529 20.5s 14:05 Araschnialevanaprorsa 1 20010529 20.5s 14:05 Canthansso. 3 20010529 20.5s 14:05 Cantharissp. 1 20010529 20.5$ 14:05 Empis ilvida 4 20010529 20.5s 14:05 Empisdiapramma 1 20010529 20.5s 14:05 Empis tessellata 3 20010529 20.5s 14:05 Enistalis arbus forum 2 20010529 20.5$ 14:05 Eristalis tenax/pertinax 1 20010529 20.5$ 14:05 Helophilusfrivittatus 1 20010529 20.5s 14:05 Het eropt era 1 20010529 20.5s 14:05 Musca sp. 3 20010529 20.5s 14:05 Scafophaqa x 1 20010529 20.5s 15:05 Araschnia levana prorsa 1 20010529 20.5s 15:05 Canthanis so. 3 20010529 20.5s 15:05 Cantharissp. 2 20010529 20.5s 15:05 Elateridae 1 20010529 20.5$ 15:05 Empis diagramma 1 20010529 20.5s 15:05 Empis tessellata 11 20010529 20.5s 15:05 Enistalis arbustorum 5 20010529 20.5$ 15:05 Erisfalis horticola 2 20010529 20.5s 15:05 Enisfalis tenax/pertinax 6 20010529 20.5$ 15:05 Helophilus fnivittatus 1 20010529 20.5s 15:05 Heteroptena 1 20010529 20.5$ 15:05 Musca sp. 3 20010529 20.5$ 15:05 Sarcophaga so. 2 20010529 20.5$ 15:05 Scat ophaqa x 2
I yyyymmdd date verge time speciesinsect 3=Lamium2=hermaphroditic album inf. species1=female tollowed (sec)time visited inf. # visited inf.# ti.on visited fI.# distances sum of flight(cm) on visitedfi.# visitedof # fi. inf. flight distancebetween inf. avg.(cm) per sec.visited avg.# (I. 20010503 19.5 13:40 Apis mellifera Apis mellifera 1 6235 25 11 8 0.73 0.13 2001050320010503 19.519.5 13:40 Apis mellifera 1 15314670 462110 1555624 612617 0.390.460.71 0.400.180.24 20010503 19.5 13:4013:35 Bombuspascuorum Apis mellifera 2 1 26713212126 241031 5 1143714 1550 9 311177112 0.410.640.44 17.7012.960.003.61 0.110.350.19 20010503 19.5 13:3513:40 Bombuspascuorum MelanostomaMelanostomasp./Platycheirussp.Bombusterrestris sp./Platycheirus sp. 2 49805363 2736 29 4 13 83 2108190 0.280.750.45 45.0035.0011.570.00 0.210.100.06 20010503 19.5 13:4013:40 RhingiacampestrisRhingia campestris 22 907052 443 11 76 43 103 0.570.270.67 25.75 0.060.08 2001050820010503 18.519.5 14:4413:40 Bombuspascuorum MelanostomaspiPlatycheinissp.Rhingia campestris 32 133239 57 379 31236121 1019 39 5569 9 0.130.310.43 18.339.86 0.050.140.04 20010508 18.518.5 14:44 RhingiaMelanostomasp./Platycheirussp. campestris 33 362267645 2 1 1514 6 32 1 15 0 0.200.330.070.32 7.503.00 0.040.03 200105082001050920010508 18.5 2 14:4411:20 RhingiaRhingiacampestris campestris 23 2731245363 17423 2437 26 53 120397015 0.210.08 30.0013.004.127.50 0.490.02 20010521 40 13:1514:1514:0014:54 BombusBombuspascuorum pascuonim 22 74676056 131814 6 43 23222527 108607357 0.51 18.004.624.064.07 0.310.330.420.48 2001052120010521 4040 14:5413:3013:45 BombusBombuspascuorum terrestns Rhingia campestris 2 1356835 26 3 1 8 41+ 3 1 165+ 20 0.131.58 0.67 0.090.01 * protocols not included on floppy no value meansnot observed FollowingAppendixI. = observationsflower; inf. N = inflorescence on Glechoma hederacea J yyyymmdd date time verge species insect visited # umb.es # visitedi # of flight sum avg. flightdistance visiting total visited # fraction umb.es visited I 20010523 15:41 36 Empis lividadiagramma umbels -12 on visitedumbels 50 umbelsi distances 8 I (cm) 15 betweenumbels 7.5 (sec)time 50 perumb.es sec. 0.160 of #visited umb.es umbels on 0.16 20010523 15:41 36 Empis tessellata 2 1 3010 14 7 65 32.5 325 34 0.2060.022 0.700.23 2001052320010529 11:5715:2715:41 20.5 36 Cantharissp.Muscasp.Canthanssp. 0 1 2010 57 4020 0 20.040.0 297300100 0.0170.0700.047 0.250.70 20010529 12:3714:3512:00 20.5 ColeopteraCantharissp.Canthanssp. rest 01 1014 9 17 48 80100 10.080.0 .460227150 0.0180.0530.037 0.400.571.89 20010529 12:1913:30 20.5 DipteraTipulidae rest (snaveMieg) 10 1410 8 126 1 0 1 1.0 17818239 0.0260.066 0.101.50 20010529 12:0013:3012:19 20.5 Empissp.EmpisDiptera livida small 1 1819 24 30 1 30.0 1.0 4275 0.0480.0530.034 0.430.110.21 20010529 12:00 20.5 Empissp.Emplssp. 1 1826 7 42 1 10040 2 100.040.0 3428 0.0710.029 0.140.08 2001052920010529 13:30 20.520.5 Empissp. 1 1315 37 40 2 40.02.0 343855 0.0880.1840.073 0.230.470.22 20010529 12:0013:30 20.5 Empissp.Empis sp.sp. 42 56222029 1311 26 3601006080 90.050.040.030.0 100235062 0.2100.0870.1100.120 0.230.090.550.21 2001052920010529 14:4514:3512:00 20.5 EmpisEmpissp. diagramma 286 29775556 3412 9 1822902042 30.348.310.05.3 30043514088 0.0300.0780.1360.064 0.310.440.220.16 20010529 11:4513:3013:4811:41 20.5 EmplsdiagrammaEmpis diagramma 435 74392740 23261212 1257431 5 25.018.510.3 1.7 120205649125 0.1000.1120.0400.096 0.160.590.960.30 20010529 11:4513:30 20.5 EnstalisEmpis diagramma a,bustonjm 0 16 79 8 27 3 75 0 12.5 35030 0.1000.077 0.380.34 20010529 12:0015:3011:45 20.5 EnstalisarbustorumEristalisEnstalis arbustorum 42 425417 9 2410128 140958030 23.840.030.0 130708060 0.1430.1500.133 0.190.710.89 following observations on Anthriscus sylvestris 20010529 12:3715:3015:2712:00 20.5 EristalishorticolaEnstalisErlstalis arbustorumarbustonim 384 367234 392213 1103566 11.713.816.535.0 16014489 0.1460.2440.1530.185 0.360.650.570.54 umb.esAppendix = umbellules 0 no value meansnot observed I 4 20010529 14:35 20.5 Eristalis tenax/pertinax 12:19 20.5 Eristalis horticola 09 1 8513 33 5 173 0 19.2 15857 0.0880.209 0.380.39 20010529 13:3012:00 20.5 EristalisEnstalis tenax/pertinax 13:48 20.5 Erlstalls tenax/pertinax 2 32313632 121058 405080 4 20.025.080.02.0 90503028 0.2860.1330.2000.167 0.330.310.160.26 20010529 14:3512:1913:30 20.520.5 EristalisEristalisErlstalisEristalistenaxlpertinax tenax/pertinaxtenaxlpertinax 43 444136 11 5 1203285 40.028.38.0 1079254 0.2040.1030.054 0.250.270.14 20010529 15:3012:0013:30 20.520.5 EristalisEristalisEnstalistenaxlpertinax tenax/pertinaxtenaxlpertinax 10 65 94687262 28552317 150548255 30.013.79.25.4 54811612373 0.1000.3150.1470.228 0.590.340.240.45 20010529 13:3014:35 20.5 HelophilusErlstallstenax/pertinax trMttatus 11 3 1 1163114 1035 3 2257550 25.050.020.5 1958026 0.1250.1150.179 0.320.210.30 20010529 11:41 13:3012:19 20.520.5 Helophilus Muscasp. trlvittatus 20.5 Muscasp. 30 1 412811 734 25 90 25.0 3.0 783576 0.0860.0530.090 0.170.110.36 20010529 12:3712:5512:00 20.520.5 SarcophagaMuscaSarcophagasp. sp. sp. 1 2115 64 10030 100.030.0 4025 0.1500.160 0.290.27 20010529 11:4111:4514:3513:30 20.5 Tachina fabnciafabrlcia 20.5 Tachina fabrlcla 272 77243427 652310 9 1021503035 75.015.017.514.6 2911343066 0.2230.3330.1720.136 0.680.330.840.42 23 20010503 18 11:17 1 Apis mellifera 01 5015 5115 0.00 65 54 20010503 18 11:2911:21 2 1 Apis mellifera 74 0 3553 127 35 0.000.580.02 67 876 20010503 18 11:5011:3711:34 1 Apis mellifera 502 8834 8836 0.000.06 67 109 20010503 18 12:0011:55 1 BombusApis melilferamellifera terrestns 1507 1 1310 2665 2817 2770 0.530.040.07 7 14131211 20010503 19.5 13:101818 12:1712:12 21 ApisApis mellifera me//iferamellifera 578600 9081 26514659 578659 236346 0.380.231.00 376 18171615 20010503 19.5 14:3014:35 22 BombusRhingia campestristerrestrispascuorum 22732920 4639 499815 1920 27723735 6559 0.780.710.66 675 2019 20010503 19.5 14:30 2 1 Apis mellifera 44 2 38 6 82 8 0.250.540.82 37 232221 20010503 19.5 14:4714:4514:40 2 1 ApisBombus mellifera lone//uspascuorum 1610546246 32211292 1932638358 0.830.860.69 876 27262524 20010503 19.5 15:0014:5014:53 2 I1 ApidaeApisBombus mellifera small pascuorum 15234 346 3519 542191 5638 15776 537 7573 0.970.640.250.48 376 302928 20010503 19.5 15:0015:1015:05 2 1 ApisBombus mellifera pascuorum 1457765 46 23721325 1694978 71 0.650.860.78 7 34333231 2001050920010503 19.5 15:122 11:5711:4911:45 2 BombusMelanostoma pascuorum spiPlatycheirus sp. 1018854274418 1763750 5 10301055324423 0.960.850.99 667 pollenGlechoma load slides hederacea Appendix P ______•1
# slide date vergetime insect # target # rest total #fraction yyyymmdd species pollen pollen pollen target ponen Appendix 0 grainsgrainsgrains pollenspecies pollen load slides 3 Anthriscus sylvestns 35 20010521 5 12:30 Musca sp. 0 12 12 0.00 4 36 20010521 5 12:30 Scatophaga stercoraria 0 9 9 0.00 no value = not observed I 37 20010523 36 Empis tessellata 2372 76 2448 0.97 9 4 38 20010523 36 Empis diagramma 2130 25 2155 0.99 5 39 20010523 36 Empis tessellata 4120 38 4158 0.99 5 40 20010523 36 Empis tessellata 1042 13 1055 0.99 0.97 6 41 20010523 36 Empis livida 1625 48 1673 42 20010523 36 Empisdiagramma 2055 26 2081 0.99 7 43 20010523 36 Empis lMda 1165 93 1258 0.93 9 10 44 20010523 36 Empis liinda 1036 115 1151 0.90 6 45 20010523 36 Gymnochaeta vindis 1219 49 1268 0.96 4620010523 36 Empis tessellata 2440 4 2444 1.00 4 47 20010523 36 1-lelophilusyellow 2560 9 2569 1.00 5 4820010523 36 Diptera small black 255 20 275 0.93 6 4920010523 36 Enstalis horticola 4841 98 4939 0.98 11 9 5020010523 36 Empis ilvida 4467 62 4529 0.99 0.99 10 51 20010523 36 Gymnochaeta vindis 2972 27 2999 52 20010523 36 Lepidoptera 669 28 697 0.96 6 5320010523 36 Empis tessellata 4946 12 4958 1.00 5 54 20010523 36 Helophilus yellow 3526 70 3596 0.98 10 6 55 20010523 36 Empis diagramma 2430 44 2474 0.98 9 56 20010523 36 Diptera small black 764 27 791 0.97 57 20010523 36 Coleoptera green 1000 73 1073 0.93 8 5820010523 36 Musca sp. 891 11 902 0.99 5 4 59 20010523 36 Dipt era 4369 9 4378 1.00 8 60 20010523 36 Empis tessellata 3143 104 3247 0.97 0.94 7 61 20010523 36 Empis livida 340 21 361 6 62 20010523 36 Eristalis sp. 515 14 529 0.97 9 6320010523 36 Symphyta 1050 20 1070 0.98 10 6420010523 36 Empis tessellata 1863 35 1898 0.98 65 20010523 36 Gymnochaeta viridis 1556 181 1737 0.90 11 11 66 20010523 36 Sarcophaga sp. 1743 30 1773 0.98 10 67 20010523 36 Empis livida 4465 137 4602 0.97 10 68 20010523 36 Gymnocha eta viridis 405 89 494 0.82 6 69 20010523 36 Musca sp. 4390 42 4432 0.99 7 7020010523 36 Gymnochaeta vindis 2890 42 2932 0.99 7 71 20010523 36 Empis tessellata 3938 42 3980 0.99 72 20010523 36 Gymnocha eta vindis 3164 88 3252 0.97 11 7320010523 36 Empis diagramma 2050 62 2112 0.97 9 7420010523 36 Empis tessellata 4608 16 4624 1.00 5 4 75 20010523 36 Scatophaga x 1726 24 1750 0.99 6 76 20010523 36 Empis tessellata 2190 34 2224 0.98 7720010523 36 Tachina fabncia 3262 26 3288 0.99 5 8 78 20010523 36 Tachina fabncia 3490 112 3602 0.97 7920010523 36 Enstalis horticola 7350 808 8158 0.90 8 9 80 20010523 36 Eristalis sp. 4112 72 4184 0.98 0.89 8 81 20010523 36 Lepidoptera 478 60 538 9 82 20010523 36 Enstalis sp. 1860 46 1906 0.98 83 20010523 36 Empis diagranjma 4440 176 4616 0.96 9 8420010523 36 Canthans sp. 2832 56 2888 0.98 4 8520010523 36 Scatophaga x 732 34 766 0.96 7 8620010523 36 Diptera small black 288 46 334 0.86 7 87 20010523 36 Diptera small black 218 10 228 0.96 6 I'
16 1104 0.99 5 88 20010523 36 Empis diagramma 1088 4894 110 5004 0.98 8 8920010523 36 Empis livida 18 874 0.98 4 90 20010523 36 1-leteroptera small 856 78 2176 0.96 10 91 20010523 36 Empis livida 2098 148 10520 0.99 11 9220010523 36 Tachina fabncia 10372 62 3168 0.98 10 9320010523 36 Gymnochaeta vindis 3106 66 2038 0.97 8 94 20010523 36 Empis diagramma 1972 42 1634 0.97 6 9520010523 36 Elateridae 1592 2692 30 2722 0.99 7 96 20010523 36 Empis diagramma 5162 0.98 12 1 50a 20010605 36 Lepidoptera 5084 78 8132 0.98 9 150b 20010605 36 Eristalis horticola 7984 148 988 0.99 3 151 a 20010605 36 Empis diagramma 980 8 1476 0.99 2 151b 20010605 36 Cantharissp. 1460 16 1196 5343 0.78 9 1 52a 20010605 36 Helophhlus black 4147 2856 0.91 5 152b 20010605 36 Empis tessellata 2600 256 112 8500 0.99 5 1 53a 20010605 36 Empis tessellata 8388 736 0.91 3 153b 20010605 36 Elatendae 668 68 1020 0.97 3 1 54a 20010605 36 Scatophaga stercorari 992 28 0.99 2 1Mb 20010605 36 Empis tessellata 1163 16 1179 11260 0.96 10 155a 20010605 36 EristalhshortiCOIa 10844 416 1012 0.96 4 155b 20010605 36 Coleoptera green 972 40 30 6070 1.00 3 1 56a 20010605 36 Scatophaga stercoraria6040 64 5720 0.99 2 I 56b 20010605 36 Enstaiis horticola 5656 2648 0.99 3 157a 20010605 36 Symphyta 2632 16 10680 1.00 5 15Th 20010605 36 Canthans sp. 10640 40 128 1.00 1 158a 20010605 36 Elateridae 128 0 904 0.95 3 158b 20010605 36 Ensfalis arbustorum 856 48 1576 64 1640 0.96 3 1 59a 20010605 36 Helophilus yellow 3352 15192 0.78 5 1 59b 20010605 36 Erisfalis horticola 11840 896 0.94 3 160a 20010605 36 Scatophagax 840 56 1056 0.97 2 160b 20010605 36 Scatophaga x 1024 32 11856 1.00 4 161a 20010605 36 Cantharissp. 11840 16 40 1184 0.97 3 161b 20010605 36 Scatophagax 1144 440 0.93 2 162a 20010605 36 Scafophaga x 408 32 64 1384 0.95 3 162b 20010605 36 Scatophaga x 1320 1360 24 1384 0.98 2 1 63a 20010605 36 Canthans sp. 48 2208 0.98 2 1 63b 20010605 36 Empis diagramma 2160 240 0.97 2 164a 20010605 36 Scat ophaga x 232 8 2424 0.99 2 1Mb 20010605 36 Scatophaga x 2392 32 96 9192 0.99 7 1 65a 20010605 36 Eristalis horticola 9096 528 3712 0.86 5 1 65b 20010605 36 Eristalis arbustonm 3184 64 896 0.93 6 166a 20010605 36 Scafophaga x 832 644 10620 0.94 8 I 66b 20010605 36 Eristalis horticola 9976 I slide vyyymmdd date time Ivergel speciesinsect I from plant type 1=female I to plant type 1=temale I numberI pistil I target I # rest # I total # I fractiontarget I pollen I # I 1 20010515 10:33 2 Bombus pascuorum 2=hermaphroditic 3=both 2 I 2=hermaphroditic 3=both 2 1 grainspollen I 26 grainspollen I pollengrains I 2 28 grainspollen I 0.93 I speciesi 2 I 324 20010515 10:33 2 BombusBombuspascuorum pascuorum 22 1 2 32 1 97 1 43 1211 5 0.200.750.64 42 65 20010515 10:39 2 Bombuspascuorum 1 22 32 1 3017 0 3017 1.00 1 1 87 20010515 10:4810:39 2 BombusterrestnsBombusterrestrisBombuspascuorum 1 21 2 120 100 120 0.17 4 11109 20010515 10:48 2 BombusterrestnsBombus terrestris 1 1 1 1 143 73 13 3 20 64 0.500.25 532 no value meansnot observed 1213 20010515 10:54 22 Bombuspascuorum 1 1 132 0 56 56 0.000.350.00 4 1514 20010515 10:54 2 BombusBombuspascuorum pascuorum 1 1 2 1 13 0 743 43 0.00 43 19181716 20010515 10:5411:24 2 BombusBombuspascuorum pascuorum 2 1 2 1 2 1 304 14 24 2018 54 0.600.000.220.65 53 2120 20010515 11:3111:24 2 RhinqlaBombuspascuorum campestris 22 1 22 3 1 42 3 1 5 0.800.40 24 2322 20010515 11:31 2 RhinqiacampestnsRhinqiacampestris 1 2 32 1 1210 4 023 127 0.570.831.00 43 1 262524 20010515 11:34 2 BombuspascuorumBombuspascuorum 2 21 132 19 5 02 21 5 1.000.90 2 1 2827 2001051520010515 11:55 2 Rhinqia campestns 2 1 2 1 024 310 34 0.671.00 2 1 302931 20010515 12:1812:1511:55 22 RhinqiacampestnsRhinqia campestns 2 1 22 2 1 101 78 196 1 274102 1 0.990.280.00 16 3332 20010515 12:1812:18 2 BombusBombuspascuorum pascuorum 1 21 32 1 290 60 89 0.001.00 1 3534 20010515 12:20 2 BombusBombuspratorum pratorum 3 1 32 1 21 5 37 2412 0.880.420.25 3 383736 20010515 12:2712:20 2 referenceRhinqiaBombuspratorum campestns hermaphroditic 1 1 079 103 1 1210 8 0.750.00 23 414039 20010515 12:47 2 RhlnqiaRhinqlareference campestnscampestris female 2 2 21 35 8 3 1 38 9 0.920.890.88 2 Appendix R 42 20010515 12:47 2 Rhinqia campestns 2 2 13 4152 3 4453 0.98 42 slides of pollen deposition 454443 20010515 12:47 2 RhinqlaRhinqiaRhinqia campestrlscampestns campestns 2 1 2 12 10 8 24 12 0.670.830.93 3 on virgin Glechoma hederacea 484746 20010515 12:5412:47 2 BombusRhinqiaRhlnqia campestnscampestrlsterrestns ? 1 2 1 32 1 302 24 546 0.000.330.60 34 pistils A 5049 20010515 12:5912:54 2 BombusBombuspascuorum terrestris ? 1 1 12 80 0 1 90 1 0.89 20 1 5251 20010515 13:0513:05 22 BombuspascuorumBombus pascuorum 1 1 2 1 0 1 34 35 0.00 32 555453 20010515 13:0713:05 2 BombusBoinbuspascuorum terrestris 3 1 3 1 2 1 0 1 02 12 0.000.201.00 12 1 5756 20010515 13:12 2 AnthochariscardaminesGorepteryxrhamni 2 21 1 100 0 1 01 100 1 1.000.00 1 58-358-258-1 20010515 13:30 Rhinçjia campestris 2 1 1 1 0 1 21 2 0.000.50 2 616059 20010515 13:3413:32 2 AnthocharisRhinqia campestris cardamines 2 1 1 1 0 320 320 0.00 20 62 20010515 13:52 2 Melanostoma sp./Platvcheirussp/Platvchein,s sp. 1 2 1 1 0 1 42 42 0.00 4 1 656463 20010515 13:52 2 Apidae small 1 2 1 2 1 1 30 4 1 0.251.00 41 6766 20010515 14:0113:58 22 Apidaesmall3Apidaesmall2Apidae small2 2 1 1 2 1 15 3 1 164 1 0.940.25 123 706968 2001051520010515 14:0114:07 2 Apidaesmall3RhinqiaApidaesmall3 campestns 21 2 1 32 1 40 0 1 81 48 2 0.830.500.00 32 737271 20010515 14:07 2 BombuspascuorumRhinqia campestriscampestns 2 2 2213 11072 1008 151211 0 0.470.171.00 570 1 767574 20010515 14:10 2 RhinqiaBombuspascuorum campestrls 2 2 1 213 082 641 12 63 0.67 342 7877 20010515 14:40 2 RhinqiaRhingia campestriscampestrls 2 1 3 1 60 23 83 0.750.00 2 13 8180 20010521 22 Apis mel/heramellifera 2 2 32 1 32 2 0 1 33 2 0.971.00 2 848382 2001052120010521 2 ApidaelargeApisMelanostoma mellifera sp./Platychei,us sp. 2 2 1 1 48 12 06 54 2 0.891.00 15 8685 20010521 2 ApidaeSyrphidae large 1 1 2 1 0 42 43 0.000.33 3 8887 20010521 2 ApidaelarqeBombuspascuorum 2 2 2 1 15 028 13 35 0.381.00 34 1 919089 20010521 2 SyrphidaeBombus pascuorumpascuotum 2 2 3 1 54 0 10 0 640 0.840.33 20 95949392 20010521 2 BombusApidae large pascuorum 2 2 321 02 4632 465 0.000.400.50 34 opP000000. Ih r1L I lIttlE! uiE!w w • tt0) I I NH 00N N NH N N NH NH NH NN 00 000000 o.u.4NNNNNNNNNN 0000 1 0 Ua 5a - a. I 0 .4 date (yyyymmdd) insect species red of red white of red orange of or. white of or. 1 1 yellow of y. white of y. 1 blank total 20010612 ApidaeAnasimyasp. large 0 0 01 0 0 1 0 41 271 2001061220010612 ApidaeDiptera small small 0 01 01 0 0 0 0 1 1 20010612 GymnochaetaEristalisEmpis sp. arbustorum/abusivus viridis 501 133 41 2 06 61 149 5017 20010612 Helophilus largesmall 1018 1 1816 2010 23 9 41 48 4726 12787 1 20010612 MelanostomaHymenoptera sp./Platycheirus(sluipwesp) sp. 24 30 02 501 205 240 1630 1039 20010612 ScatophagastercorariaSarcophagasp.Musca sp. 0 0 21 23 02 0 4 1 78 20010612 SyrphidaeScatophagax (bijzweef) 0 10 0 01 0 0 021 21 20010612 Syrphidae(doodshzw.)Tachinafabricia 01 0 0 0 0 0 1 365 2 Discrimination experiment with fluorescent dye powder on Aegopodium podagraria. Appendix U or.WhiteHalfThe = total colouredorange, of redobservation means yumbels = yellow the timewere uncoloured was presented. 1.5 hours. half of the red coloured umbel _____
Appendix V weather conditions during observation days (estimated)
Idate Itime Itemperature_(°C)Iwind__Isky I 2001050810:45-11:08 13 cloudy/sunny 2001050814:44-15:12 16 starts 2001050914:49 19 very 2001050910:24 16 yes sunny 2001050910:55 16 very 2001050912:22 17 yes sunny 200105109:35-11:18 16 yes 2001051011:38-12:56 20 yes sunny 2001051013:57-14:37 24 yes sunny 2001051014:50 24 yes 2001060510:50-11:30 18 yes 2001060512:00-12:40 17 cloudy 2001052113:15-1 4:37 12+ cloudy and sunny 2001052114:54 15-i- cloudy Appendix W List of figures and tables Page
8 Figure 1. Research area 13 Figure 2. Breeding system of Glechoma hederacea 14 Figure 3. Seed set Glechoma hederacea in the field 15 Figure 4. Nectar production by Glechoma hederacea 16 Figure 5. Nectar production by Lamium album 17 Figure 6. Sugar concentration of Lamium album nectar 18 Figure 7. Fraction of visited Glechoma hederacea visitors 20 Figure 8. Visitors on Glechoma hederacea 21 Figure 9. Visitors on Glechoma hederacea in three different areas 22 Figure 10. Flight distances of Glechoma hederacea visitors 24 Figure 11. Visitation speed of Glechoma hederacea visitors, data from following observations 25 Figure 12. Visitation speed of Glechoma hederacea visitors, data from plot observations 26 Figure 13. Percentage visited flowers per Glechoma hederacea inflorescence, data from following observations 27 Figure 14. Percentage visited flowers per Glechoma hederacea inflorescence, data from plot observations 28 Figure 15. Pollen loads of Glechoma hederacea visitors 30 Figure 16. Pollen deposition on virgin Glechoma hederacea pistils 31 Figure 17. Pollen species on Glechoma hederacea pistils 32/33 Figure 1 8a. Red fluorescent dye powder on Glechoma hederacea 32/33 Figure 1 8b. Yellow fluorescent dye powder on Glechoma hederacea 32/33 Figure 1 8c. Green fluorescent dye powder on Glechoma hederacea 34 Figure 19. Breeding system of Anthnscus sylvestris 36 Figure 20. Relative abundance of the main Anthnscus sy!vestns visitors 37 Figure 21. Visitors on Anthriscus sy!vestns 38 Figure 22. Visitors on Aegopodium podagraria 39 Figure 23. Flight distances of Anthriscus sylvestns visitors 41 Figure 24. Visitation speed of Anthriscus sylvestris visitors 42 Figure 25. Percentage visited umbellules per Anthriscus sylvestris umbel 43 Figure 26. Pollen loads of Anthriscus sylvestris visitors 44/45 Figure 27a. Red fluorescent dye powder on Anthriscus sylvestris 44/45 Figure 27b. Yellow fluorescent dye powder on Anthriscus sylvestris 44/45 Figure 27c. Green fluorescent dye powder on Anthriscus sylvestris 46 Figure 28. Visitors on Aegopodium podagraria umbels with fluorescent dye powder 47 Figure 29. Fraction of insect visitors with different specialisation levels 50 Figure 30. Visitor guild composition Glechoma hederacea Ap. B Figure 31. Verge visitor composition 2000 Ap. C Figure 32. Nectar production by Glechoma hederacea, divided into plant types Ap. D Figure 33. Pollen grains on Glechoma hederacea virgin pistils 12 Table 1. Flowers in verges 23 Table 2. Maximum flight distance between inflorescences of Glechoma hederacea visitors 29 Table 3. Characteristics pollen loads of Glechoma hederacea visitors 40 Table 4. Maximum flight distance between umbels of Ant hriscus sylvestns visitors 44 Table 5. Fraction Anthriscus sylvestns pollen in the pollen loads of Anthriscus sylvestris visitors. 49 Table 6. An overview of traits of visiting insect species on Glechoma hederacea 53 Table 7a. An overview of indexed effectiveness values for Glechoma hederacea female visitors 53 Table 7b. An overview of indexed effectiveness values for Glechoma hederacea hermaphroditic visitors 54 Table 8. An overview of traits of visiting insect species on Anthriscus syl vest ris 56 Table 9. An overview of indexed effectiveness values for Ant hriscus sylvestris visitors Appendix X Glechoma hederacea
700x
Hondsdraf (Glechoma hederacea) A bloemstengel met tweeslachtige boemen; B bloemstengelmet vrouwelijke bloemen; C vrouwehjke bloem; D tweesachtige bloem; E meeldraden; F stuifmeelkorrel (stephanocolpaat):1 polair,2equatoriaal, 3 korrelopperviak; G stamper; H vruchtbeginsel met nectarium; I stempels;J nootje.
maandblad voor imkers juli/augustus 2001 .o a pCD Appendix Z Thesis description
Pollination effectiveness of insect visitors on Lamium album, Glechoma hederacea (Lamiaceae) and Anthriscus sylvestris (U mbel I iferae) Research Proposal by Maaike de VIas, March-September 2001 Supervisors: Manja Kwak & Frank Hotfmann, Laboratory of Plant Ecology
Introduction In Northwest Europe, 80% of the flora is pollinated by insects. This part of the flora is said to be entomophilous (Kwak, 1994a). Mostly a number of insect species acts as the functional group of pollinators, called the pollinator guild. Some important insect species could fall out in the case of an environmental change (Kwak, 1994a). In restoration ecology it is important to predict possible reestablishment in presence of the available pollinators. Therefore, more information is needed on the relative importance of insect species in existing plant populations. In some plant communities, pollinator keystone species could be assigned (Memmott, 1999). Memmott (1999) has proposed a prototype foodweb. This foodweb is based on insect visitation. If the pollination effectiveness of the different insect species is introduced as a factor into this visitation web, a real pollination web can be made. Insects classified as keystone species in the food web may play a second fiddle as pollinator. Olsen (1997) defines pollinator effectiveness as the number of seeds that are 'produced' after a single visit. This depends on the number of conspecific pollen grains individuals of a particular insect species transfer (Herrera, 1987). A small population often has to cope with a heterogenous pollen deposition, because the bulk of the surrounding flowers are of another species (Kwak et al., 1998), competing for pollinators (l-leinrich, 1979). Pollinator importance is defined as: pollinator effectiveness x relative abundance of visits.
The main question is, whether visitation and pollination by insects of target plant species differ in areas that differ in plant and insect species richness. The main question can be divided into three sub-questions that will be investigated in three different areas: 1.Which insect species and how many of them visit the three plant species? 2.Which insect species does effectively pollinate those plant species? 3. Do pollinators differ in flower constancy and flight distance and what are the consequences for pollination?
Materials and methods In this study, Lamium album, Glechoma hederacea (Lamiaceae) and Anthnscus sy!vesfris(Umbelliferae)were chosen as study species. In The Netherlands, these species are often seen in road verges. Road verges are mown several times per year. This mowing regime can influence the number of flowers at a time and the microhabitat necessary for pollinating insects. Lamium album and Glechomahederacea(Lamiaceae) are both visited (and pollinated) by bumblebees (Proctor et al., 1996). Important visitors of Anthnscus sylvestns belong to Diptera: Scatophagidae and Muscidae (Hoffmann, unpublished data). Figures la, lb and ic (Hoffmann, unpublished data) show the insect species or insect groups that could be expected on the plant species in three different areas. Two or three areas with a difference in plant and insect species richness will be chosen as research areas. Size of patches and density of flowers of the three plant species will be measured by counting flower heads or umbels.
1. Visiting insect species Individuals on the Lamiaceae will be followed in plots. in plots, the fate of individual insects can be followed. Detailed information about number of visits per species will be obtained. Furthermore visitation rate can and will be measured. The flowers of Anthriscus sylvestns are too small and are visited by too many insects to be able to do proper plot observations. In transects, the overall composition of the guild can be estimated (Dramstad, 1996). Visitation of the three plant species will be followed in transects. The quick overview gives the number of individuals per species. Transect observations will be made for the three plant species. The data obtained by transect obse,vations could be presented as foodweb, in the way proposed by Memmott (1999). The data obtained by plot obseivations could be presented in a table, containing insect species, number of visits, time per visit and average flight distance.
2. Effectively pollinating Insect species Of some flowers which are virgin, the insect species have to be counted and subsequent seed set measured. To prevent visitation in larger flowers, umbels or wholeplants, net bags will be used. Deposited pollen will be counted and identified microscopically onstigmas that have been visited. ReceptMty of the visited stigmas will be determined by comparingtheir morphology to the morphology of a control set that has been stained or tested for esterase presence(Dafni, 1992). The results could be presented in a table. The rows are insect species; columns are number of pollen deposited, pollen species, receptivity of the stigma and seed set of the plant species. Results of visitation and effectiveness can be combined, to produce pollination by an insect species: pollination success (seed set) x number of visits. Crossing experiments have to be done, so that something can be said about the overall effectivity of pollination by insects. Seed set of flowers that are open (insect) pollinated will be compared to seed set of flowers that are extra hand cross-pollinated. Hand-pollination is a kind of control on resource availability (Widen & Widen, 1990) when compared to natural insect pollination. Hand cross-pollination will be done by putting an anther against a stigma (Corbet, 1999) or by using a brush. From the hand cross-pollination, maximum seed set per fruit can be calculated (Herrera, 1987). The plants will be netted and kept in an experimental garden to prevent loosing the plants by means of the mowing regime and to let them develop seedsunder equal conditions. Results could be presented in a bar graph. On the y-axis is the number of seeds, on the x-axis the plant species, in their different areas. The bars can be divided according tothe way the plant has been pollinated (normal or extra).
3. Flight distance and flower constancy The choices of the bumblebee species as a whole are defined by the choices of individual bumblebees, and bumblebee workers can have temporal flower constancy (Kwak & Tieleman, 1994c). Therefore, it is interesting to mark (and follow) bumblebees in the area, which makes it also possible to analyze flown distances between two resights of the sameindividual. Marking can be done in different ways (Walther-Hellwig & Frankl, 2000a; Dramstad, 1996; Heinrich, 1979), but the method described in Kwak (1987) seems to be most convenient.With only five colours, 499 insects can be clearly marked. The distance of the pollen transfer can be estimated by applying fluorescent dye powder at different plants (Kwak, 1994b) and collecting the fluorescent dye colors. Itis also possible to spot transferred powder from a plant on surrounding plants. Results can be presented in a scatter plot, with distance on the y-axis and plant species on the x-axis. Another graph can show the number of fluorescentdye powder particles found at different distances. Beattie (1972) describes how to remove and stain pollen from insects: the body load can be removed with a sticky staining gel (Kwak & Velterop, 1997). To be able toidentify pollen loads on insects, pollen from surrounding flowering plants have to becollected (Kearns & lnouye, 1993). Results could be presented in a bar graph. Number of pollen is on the y-axis, insect species on the x-axis. The bars can be divided into pollen species. References
Beanie,A.J., 1972.A technique forthestudy of insect-borne pollen. Pan-Pacif Entomologist 47:82 Cof'bet, S.H., 1999. Spatiotemporal patterns in the flowering of bluebell, Hyacinthoides non-scnpta(Hyacinthaceae).Flora 194,345-356 Dafnl, A., 1992. Pollination Ecology - A practical Approach. The Practical Approach Series; Series editors:D.Rickwood and B.D. Names: Oxford University Press Dramstad, W.E., 1996. Do bumblebees (Hymenoptera:Apidae) Really Forage Close to Their Nests? J. of Ins. Beb. 9(2): 163-182 Helnrlch, B., 1979. Bumblebee economics—Harvard University Press, Cambridge, Massachusetts, and London, England Herrera, CM., 1987. Components of pollinator 'quality' : comparative analysis of a diverse insect assemblage. Olkos 50:79-90 Keams, CA. and lnouye, D.W., 1993. Techniques for Pollination Biologists — University Press of Colorado Kwak, N.M., 1987. Marking a bumblebee without anaesthesia BeeWorld 68: 180-181 Kwak, N.M., 1994a. Populatie structuur & bestuM. effecten van ruimtelijkerangschlkking bij de Zwartblauwe Rapunzel. Landschap 11(1): 15-24 Kwak, M.M., 1994b. Planten en bestuivers: achteruitgang leidt tot verschuivende relaties. Landschap 11(1): 29-39 Kwak, MM. and Tieleman, I, 1994c. Het Hommelleven — Stichting Jeugdbondsuitgeverij en Stichting Uitgeverij Koninklijke Nederlandse Natuurhistorische Vereniging. Utrecht Kwak, N.M. and Velterop, 0., 1997. Flower visitation by generalists and specialists: analysis of pollinator quality. Proc. Exper. & AppI. Entomoif. 8: 85-89 Kwak, M.M., Velterop, 0. and Van Andel, J., 1998. Pollen and gene flow in frageented habitats. App!. Veg. Sc!. 1:37. 54 Memmott, J., 1999. The structure of a plant.pollinator food web. Ecology Letters 2: 276-280 Olsen, K.M., 1997. Pollination effectiveness and pollinator importance in a population of Hetherotheca subaxillaris (Asteraceae). Oecologis 109: 114-121 Proctor, M., Yeo, P. and Lack, A., 1996. The Natural History of Pollination — The New Naturalist — HarperCollins Publishers Walther-Hellwlg, K. and Frankl, R., 2000a. Foraging Distances of Bombus muscon.im. Bombus lapidanus, and Bombus terrestns (Hymenoptera, Apidae). Journal of Insect Behavior, 13(2): 239-246 Widen, B. and Widen, N., 1990. Pollen limitation and distance-dependent fecundity in females of the clonal gynodioecious herb Glechorna hederacea (Larniaceae). Oecologia 83: 191-196
Research schedule (22 weeks)
March and 11t week April • Read • Write research proposal 2,d, and4thweek April • Start breeding systems: caging • Plot observations Lamiaceae • Follow bumblebees • Lamiaceae seed set in one visit May (5 weeks) • Breeding systems: hand pollination • Transect observations • Anthriscus seed set in one visit • Anthriscus fluorescent dye powder and 2' week ,June • Seed set Lamiaceae • Pollen load counting • Insect determination 3 and 4th week June, lit week July • Seed set Ant hriscus • Collect results 2d, 3'dand4thweek July and week August • Collect results • Write Figure la,lb and ic. Insect species visiting Lan,lum album - Visiting insect species composition on Lamium album (la), Glechoma hederacea(ib) and Anthriscus syWestns (lc) in •0Is0 Dipisos the three research areas. ._,_ cn!20fls Data for L. album and for G. hederaceahavebeen • eomcsa p1110W11 collected in May and June . • 8ombia II,?,sf, 2000. For those data, all U 1 •Somkm pUOflll1 verges where the plant •$omt,a ?,0vflM,, species occurred were used. Data for A. sylvestris have been collocted in the fourth week of May 2000. For those data, three verges per area were used. Ar.a Figure 1 a
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Insect speciesvisiting Anthriscussylvestris
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