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Pollination Biology of Heracleum Sphondylium L. (Apiaceae). the Advantages of Being White and Compact

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Pollination Biology of Heracleum Sphondylium L. (Apiaceae). the Advantages of Being White and Compact Vol.7l. No.2: 163-170.2002 ACTA SOCIETATISBOTANICORUM POLONIAE 163 POLLINATION BIOLOGY OF HERACLEUMSPHONDYLIUM L. (APIACEAE). THE ADVANTAGES OF BEING WHITE AND COMPACT MeRcnI ZYCH WarsawUniversity Botanic Garden Aleje Ujazdowskie4,00-478 Warszawa, Poland e-mail:[email protected]. pl (Received:August l,zmL. Accepted:February Il,2OO2) ABSTRACT Two questions were addressedin the present study: ( I ) What are the main pollinators of the two subspeciesof H. sphondylium?, and (2) Do the studied plants sharethe pollinators' set or are they attractive for different groups of insects?The survey showed that among 40 insect taxa visiting both subspeciesof H. sphondylium approx. only 53Vo caried significant pollen loads. However, the Pollinator Importance Coefficient (IC) calculated for each insect group, and based on observation of insects' abundance,within-umbel activity and pollen load revealed that only two taxa in case of H. s. ssp.sibiicum (Thicops nigifrons, Eiozona syrphoides) and four in case of H. s. ssp.sphondylium (T. nigifrons, E. syrphoides,Meliscaeva cinctella and Arge ustulata) were truly important polli- nators. Although both subspecieswere visited by similar insects, H. s. ssp. sphondylium, with its characteristic compact and white umbels, was visited more frequently by Diptera and Hymenoptera, while yellow-greenish loose umbels of H. s. ssp,sibiricum were preferred by Coleoptera.This paper indicates that the concept of faithful pollinators may also apply to a broader spectrum of Apiaceae,usually consideredprimitive in terms of pollination strategies,and suggestspossible ways of differentiation in two closely related taxa. KEY WORDS: Heracleum sphondylium L., Diptera, colour morphs, pollination, pollinator importance. INTRODUCTION tegazzianum(Grace and Nelson 1981) showed a considera- ble difference in pollinators' importance. Analysis of in- For the last decade the traditional view of plant pollina- sect visitors of two American umbelliferean genera Tha- tion systemshas been questionedwith arising evidencefor spium and Zizia showed that for most pollinations (up to widespread generalisation of plant pollination strategies 75Vo) orly few insect species were responsible (Lindsey and opinion that the previously accepted idea of a direct 1984). It is also noteworthy that in case of closely related link between the observed floral traits and pollinators mi- H. sphondylium and H. mantegazzianum,growing sympa- ght not be so simple (Waser et al. 1996; Johnson and Stei- trically in a mixed population and characterisedby similar ner 2000). As shown by different authors, plants conside- pollination systems,different pollinators' assemblageswe- red specialised may be pollinated by a considerablenum- re observed (Grace and Nelson 1981). It may have been ber of taxa (Beattie 1971), and primitive species,as those caused by the difference in heights of the inflorescence. 'faithful of Apiaceae, could have pollinators' (Bell 1971). A similar situation may probably be found in other closely The descriptions of flower visitors of Apiaceae list nu- related taxa. merous insectsbelonging from severalto a few hundred ta- Apart of inflorescenceheight, flower colour and symme- xa (Drabble and Drabble 1917, 1927; Corbet 1970; Bell try may be responsible for attracting different groups of 1971; Grace and Nelson 1981; Lindsey 1984; Sheppard pollinators (Faegri and van der Pijl 1979; Proctor, Yeo and 1991; Proctor, Yeo and Lack 1996; Dlussky 1998; Lam- Lack 1996). Usually this kind of studies are conducted on bom and Ollerton 2000), only for the dipteran family Ta- the level of genera, species or for artificial flowers (e.g. chinidae Karczewski (196'7) describes33 speciesobserved Lunau 1993; Johnson and Dafni 1998). There are very few on flowers of H. sphondylium. works that focus on the consequencesof intraspecific dif- More recent studies however reveal that flower visitors ferences on the successof individuals (Herrera 1996). are not synonymous with pollinators. For example pollen Such relationship was found for two colour morphs of loads and number of visits to different speciesof Angelica Raphanus raphanistrum (Brassicaceae).In a mixed white- (Bell and Lindsey 1978) and H. sphondylium and H. man' and yellow-flowered population, individuals with yellow POLLINATION OF HERACLEU M SPHONDYLIUM ZychM. flowers were found more attractive for butterfly and hover- Study site fly pollinators (Kay 1976). Also evidence for selection of The observations were conducted in two sites in the flower colour was presented in Delphinium nelsonii same forest complex in Wigry National Park (NE Poland), (Waser and Price 1983). Further analysis of closely related the distance between the two populations was approx. 3 taxa with different flower morphology could be useful for km. The population of 11.s. ssp. sphondylium was situated tracing the possible ways of evolution of features connect- near the village Krzywe in the forest clearance approx. 50 ed with sexual reproduction and subsequentlyto show the m from the road Suwalki-Sejny (road no. 660), and the existing trends of speciesdifferentiation (Herrera 1996). So population of H. s. ssp,sibiicum on the border of the for- far no evidence on this matter has been presented for est sections 133 and 134. Both grew in similar habitat con- Apiaceae. ditions, along the forest road in mixed spruce-pine forest, Among European species Heracleum sphondylium L. and consistedof similar number of individuals. (hogweed) may serve as possible object for similar studies. Two of its subspecies(f1. s. ssp. sphondylium and ly'. s. ssp. Field observations sibiicum), formerly considered separatespecies, differ in Field observationswere conductedon23-21 July of 1998. flower and inflorescence morphology. The former has a These four days representedthe peak of flowering for the white corolla with flowers with elongated petals arranged primary umbels, also the weather was good enough to make on the edge of an umbel, while the later has greenishpetals the observationspossible. Only primary umbels with at le- 'edge with no such effect' (Gawlowska 1961, Sheppard ast50Vo of flowers in the male phasewere chosenfor obse- 1991).It seemspossible that thesefloral characteristicsmay rvations. This restriction was implied by the fact that main- serye as attractants for different groups of pollinators which ly these umbels are responsiblefor seedproduction (Shep- would create a mechanism of differentiation between the pard 1991). The observation started at 8.30 a.m. (the first two related taxa. recording) and ended at 7.00 p.m. (the last capturing). The In the present study I was especially interested in time was chosen on the basis of earlier observationsrevea- answering the following questions: ling that before 8.00 a.m. and after 7.00 p.m. flowers of ho- (1) What are the main pollinators of the two subspecies gweed are visited by insectsonly occasionally. of H. sphondylium? Every day eleven rounds of observationswere conducted (2) Do the plants studied share the pollinators' set or are (except 24rh July - nine rounds, observations stopped by they rather attractive for different groups of insects? rain; and 25th- two rounds). Each round lasted t hour and comprised of three phases: random choice of an umbel, video recording, and insect capturing. MATERIAL AND METHODS Once selectedumbels were not excluded from the next round and therefore it was possible that the same umbel H eracl eum sphondylium L. was observedtwice or more times. H. sphondylium is a perennial (hemicryptophyte) (Rabot- After an umbel had been selected it was stick bound to nov 1956, Sheppard 1991), not a biennial as often stated prevent it from wind-causedmovements. Then for 10 min- (e.g. Brummitt 1968), with cauline leaves arrangedin a ro- utes insect activities were recorded using an 8 mm video sette producing erect flower stems up to 3 m tall. It is wi- camera (Sony TR-3200). The camera was set on a tripod at despread in forests and woodland clearings, riverbanks, a distance of about 0.5 m from the umbel. The recording and tall montane-herb grasslands(Sheppard 1991). In Po- field covered the total umbel surface. 20 minutes after land it is usually described as a common meadow species recording, for l0 minutes, all the insects visiting an umbel (Gawlowska i956), growing also in thickets, on roadsides were collcted either with enthomological net (Diptera, and forest fringes (Rutkowski 1998). Its geographicalran- Hymenoptera, Lepidoptera) or directly to plastic vials ge includes most of the European countries, except the (Coleoptera,Hemiptera). Insects were killed with ethyl extreme north, parts of the Mediteranean region and some acelate and stored dry for further investigations of their Atlantic and Mediterranean islands (Brummitt 1968; Shep- pollen loads. Aphids and other small, sap sucking insects pard 1991). (e.g. Thysanoptera)were excluded from the analysis. H. sphondylium rcproduces mainly by seeds;vegetative division of root stock, although occasionally occurring, is Pollen loads analysis not important (Sheppard 1991). The open, protoandrous For investigation of pollen loads carried by insects visit- flowers, arrangedin compound umbels, are antomophilous ing flowers of H. sphondylium amodified method of Grace with insects visiting flowers for pollen and nectar excluded and Nelson (1981) and Lidsey (1984) was used.An insect by exposed swollen base of the style called stylopodium. was put on the microscopic slide and pollen from its body Most of the flowers are actinomorphic,
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