Mechanisms of Plant Evolution Special topic: speciation in Lamiaceae Special topic: speciation in the Lamiaceae Dracocephalum (Dragonhead) – a genus of 35 almost identical species The Lamiaceae challenge •Lamiaceae are around 7000 species that are often distinct in secondary compounds •Many species of Lamiaceae are crossfertile and hybrids are very common •Morphology is often very similar, taxonomy is very difficult and still unresolved („Lamiaceae are the disgrace of plant taxonomy“) What is a species in this context? Special topic: speciation in the Lamiaceae One postman, many letters – Salvia pollination •Lamiaceae are very species rich. Many species occur sympatrically. How to prevent hybridisation? •Lamiaceae flowers are zygomorph and rich in scents and patterns. They are pollinated by specialised and intelligent Hymenoptera as Bombus spec. (bumblebee). •Is speciation in the Lamiaceae maintained by specific pollinators? Pollination in Salvia. A lever mechanism deposits pollen onto the abdomen of the pollinator. The style grows later. In the receiving flower, the style takes up the pollen from the site, where it was deposited in the donor flower. Length of anthers and stamina are genetically linked. Special topic: speciation in the Lamiaceae One postman, many letters – Salvia pollination Burgenland Niederösterreich S. pratensis S. austriaca S. pratensis Conclusion: no propagation barreer by temporal difference S. verticillata •Pollinators are the same! Conclusion: no propagation barreer by specific pollinators S. nemorosa S. aethiopis •Flowering time of different species overlaps! S. glutinosa Sympatric species of Salvia. In Austria 6 species occur in the same region. Flowering times and pollinators overlap. Gene flow across species boundaries must be maintained by different mechanisms. •Due to close relationship all Salvia species should hybridise, such that there is no speciation barreer •How is speciation maintained? Special topic: speciation in the Lamiaceae One postman, many letters – Salvia pollination 4 3 1 5 6 Burgenland: 1 – S. pratensis 2 – S. austriaca 3 – S. nemorosa 4 – S. aethiopis 2 Bombus terrestris 3 14 Niederösterreich: 1 – S. pratensis 5 – S. verticillata 6 – S. glutinosa Multipurpose-Pollinator. Sympatric species of Salvia deposit pollen at different sites of the abdomen. Since length of stylum and anthers are genetically coupled, the stylum will pick up the pollen from the right site. This creates a propagation barreer. •Stamina geometry is species specific as well as length of stylum. •For each species of Salvia the stylum touches the site, where the pollen had been deposited in the donor flower. •A pollinator (such as Bombus terrestris) can transmit pollen from different Salvia species, but at different sites of the abdomen •The stylum takes up the „right“ pollen, because it is at the „right“ site. Special topic: speciation in the Lamiaceae Morphology precedes genetics: the Dragonhead case D. ruyschiana •small changes in flower morphology generate different mechanism for pollination specificity •this will lead to reproductive channelling of subclades model of pollination in D. ruyschiana (M. Sci. Stefanie Calin) D. ruyschiana D. moldavica Flower of D. moldavica (Dipl. Julia Völker) Phylogenetic tree of Dracocephalum (Ph.D. Thomas Horn)
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