Lilium Martagon

Lack oflight apossible threat toan historical Linnaean population of Lilium martagon Alexandra Holmgren Degree project inbiology, Master ofscience (1year), 2008 Examensarbete ibiologi 30 hp tillmagisterexamen, 2008 Biology Education Centre Supervisors: Mariette Manktelow and Brita Svensson Table of Contents Abstract p. 1 Introduction p. 2 Hortus Upsaliensis p. 2 Linnaeus’ Hammarby p. 2 The park p. 2 Material and methods p. 4 The location and abiotic factors p. 4 Lilium martagon L. p. 4 Myrrhis odorata (L.) Scop. p. 4 Mercurialis perennis L. p. 5 Permanent quadrats p. 5 Size measurements and qualitative inventory of L. martagon p. 5 Light measurements p. 6 Cover of M. odorata, M. perennis and trees p. 6 Removal of biomass p. 7 Unshaded control in the park p. 7 Pathogens p. 7 Self-pollination p. 7 Pollinator p. 7 Data analyses p. 8 Results p. 8 Weather conditions p. 8 Qualitative inventory p. 8 The species compared over time p. 9 Results of daylight measurements p. 9 Cover of Mecurialis perennis p. 10 The tree cover p. 10 Species and size composition in unshaded compared with shaded quadrats p. 10 Size measurements and flowers per raceme of L. martagon p. 11 Removal of shading biomass p. 11 Pathogens p. 12 Self-fertilisation test p. 12 Pollination p. 12 Discussion p. 13 Conclusions p. 15 Acknowledgements p. 16 References p. 17 Map of Hammarby Appendix A Table: Measurement and numbers of L. martagon Appendix B Table: cover of trees, M. odorata and M. perennis Appendix C Table: Actual and relative accessible light Appendix D Pathogen report: CSL York Appendix E Pathogen report: Svenska växtskyddslaboratoriet, Gothenburg Appendix F This I heard one day, listening to the casual visitors of the garden: -That Sweet Cicely is a fern, -Martagon Lily an orchid, and -Dog’s Mercury a Ground-elder Alexandra Holmgren, June 29th 2006, Hammarby Abstract The aim of this study was to find why the historical population of Martagon Lilies or Turk’s-cap Lilies Lilium martagon L. is declining at Linnaeus’s Hammarby, Uppland, Sweden. I initially proposed that they were subject to light-competition from Sweet Cicely, Myrrhis odorata, but my preliminary study from 2005 revealed that Dog’s Mercury Mercurialis perennis L. was present everywhere in the park. That species was thus introduced into the study as another possible competitor for the light. Since a large number of L. martagon exhibited shaded spots and necrosis leading to wilting, samples were examined for possible pathogens. The results revealed M. odorata to be the main competitor for light, but also that there were pathogens present in the ground and in leaves of L. martagon. My studies also revealed that the lack of light in the shaded park affected the number of L. martagon directly by leading to fewer flowers on the plants. Also, I can conclude that L. martagon growth is favoured by part light, part shade conditions. 2 Introduction Hortus Upsaliensis The internationally famous botanist and professor of medicine Carl Linnaeus worked and lived in Uppsala, Sweden. His global flora, Species Plantarum, (Linnaeus 1753) became the starting point of modern taxonomy. As a base for his studies he used the plants in Hortus Upsaliensis, the botanical garden of Uppsala. These plants eventually became type specimens for a large number of species. He introduced the still used binominal system for naming plants and animals, and the garden was arranged following the sexual system, his revolutionary way of classifying plants. Around 1800 the botanical garden was moved to its present location, the former Uppsala royal garden. At this point most of the living Linnaean heritage was lost, as one did not keep track of the 18th century plant material. The old botanical garden was reconstructed in the 1920’s, and is today known as the “Linnaeus garden”. Being a reconstruction, there is almost no plant material left there from the Linnaean era. Linnaeus’s Hammarby Linnaeus bought the estate of Hammarby in 1758 as a place where he could spend the summers with his family as well as tutor his students. For the botanical lectures he needed a minor botanical garden, and he introduced some species from Hortus Upsaliensis. We know from historical sources that he planted specimens of Martagon Lily or Turk’s-cap Lily Lilium martagon L. in front of the main building at Hammarby in 1765 (Manktelow 2001). The garden at Hammarby is nowadays a historical mirror of the botanical garden in Uppsala, as around 40 of the species Linnaeus introduced from there have survived. These populations of historical species, commonly called Linnaean species, have a historical as well as a scientific value. The property of Hammarby is today state-owned through the National Property Board, but managed by Uppsala University with co-financing from Uppsala municipality. The park The population of L. martagon is mainly situated in an area called "the park". This used to be the Linnaeus garden at Hammarby and it stretches over the eastern part of the property. The garden was an open area where Linnaeus planted exotic species and several trees (Manktelow, 2001). After Linnaeus’s death in 1778 his widow Sara Lisa Linnaea tended the garden for the remainder of her life, and then it became slowly overgrown. Linnaeus’s great-great-grandson Mauritz Ridderbjelke turned the overgrown garden into parkland, a seemingly wild, lush and romantic grove. He altered the structure of the area by creating new, meandering paths in the garden. The park became denser in trees during the 20th century. Today, most trees are approximately 50 years or younger. 3 Fig 1, Lilium martagon in the park of Hammarby on June 29th 2006. Photograph: Alexandra Holmgren The curator of Hammarby since 35 years, Kristofer de Korostenski, claims that the population of L. martagon at Hammarby is declining. He judges that is has declined with 75% in 25 years, more rapidly the last few years. During the same interval, the Linnaean species Sweet Cicely Myrrhis odorata (L.) Scop. has increased in the same area (Kristofer de Korostenski, pers. comm. 2006). A codependence between these two species was proposed by the curator, a theory supported by Mariette Manktelow, a scientist who has been studying the area for 30 years (Mariette Manktelow, pers. comm. 2006). During my own pre-study I noticed that another Linnaean species, Dog’s Mercury Mercurialis perennis L., was abundant and may be involved in the competition affecting L. martagon. A similar decline of L. martagon is known also from other places in Sweden. Large historical populations have been reported to be declining for example in the two estates Eka and Husby Gård in Uppland, Sweden (Göran Ulväng, pers. comm. 2007). Wilting of L. martagon at Hammarby was noted already by Rutger Sernander, inspector of Hammarby and professor of Botany at Uppsala University, in the 1940's (Mariette Manktelow, Karin Martinsson, pers. comm.). The purpose of my study was to investigate if light-competition in the park at Linnaeus’s Hammarby is a key factor behind the decline of L. martagon and if the abundant populations of M. odorata and M. perennis have any influence. I investigated the following two hypotheses and predictions: Hypothesis 1: The abundance of L. martagon varies inversely with the cover of M. odorata in the park. Prediction 1: Frame quadrats with high coverage of M. odorata will contain few L. martagon, and vice versa Hypothesis 2: The abundance of L. martagon varies inversely with the cover of M. perennis in the park. Prediction 2: Frame quadrats with high coverage of M. perennis will contain few L. martagon, and vice versa. Other possible factors were also investigated in the study, such as reproductive strategies, nitrogen resources and pathogens. 4 Material and Methods The location and abiotic factors Linnaeus’s Hammarby is located about 15 km SE of Uppsala, in the parish of Danmark in the county of Uppland, Sweden. The soil of the Uppsala plain consists of heavy, alkaline clay. Mean climate values 1961-1990 were: Yearly precipitation 600 mm, mean cloudiness 60%, July precipitation 60 mm, August precipitation 70 mm, sunny days in July (cloudiness less than 25% measured three times a day) 5-6. The nitrogen deposition over Uppsala is 3-6 kg/ha/year over Picea abies forests (www.sna.se) For the purpose of this study I observed the weather conditions during the summers of 2005-2007. The most common tree species in the park at Hammarby is Norway Maple Acer platanoides L., a species easily dispersed. The remnant Linnaean plant species in the ground cover are those that flower early in spring before leafing, or those that can withstand shade. The most common species covering the ground of the park were M. perennis and M. odorata. The Wood meadow-grass Poa nemoralis L. is the most common graminoid. Lilium martagon L. Martagon Lily, Turk's-cap Lily. Family: Liliaceae. Lilium martagon originates from central-eastern Europe and Siberia where it grows in mountain forests. It prefers a certain declination and a good drainage. It is considered tolerant to shade (Lundquist 2005). Growth rate is slow, in its second year it has only up to three leaves and the plant does not flower until it is five to seven years old, maybe depending on if it originated from a seed or a scale from a bulb (Kjell Lundquist, pers comm.). It reaches a height between 0.4–1.5 meter (Mossberg 2003). The population of L. martagon at Hammarby exhibits an unusually high morphological and genetic variation, which is interpreted as it being composed of several clones, as opposed to many other populations in Sweden (Lundquist 2005, Persson et al., 1998).

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