Wood anemone dispersal

My knowledge of wood anemones (Anemone nemorosa) had been gleaned from general books on woodlands and wildflower guides, effectively summed up by this extract from the National Biodiversity Network atlas:

"Wood anemones are one of the first spring flowers, their cheerful white stars appearing in March and April. They are perennials and, as their seed is rarely viable in Britain, they spread by means of underground roots. However, they do not advance their carpet quickly, and colonies of anemones can often be found in the same spot within a wood, century after century."  

https://species.nbnatlas.org/species/NBNSYS0000002700

https://commons.wikimedia.org/wiki/File:170_Anemone_nemorosa.jpg#file

The above illustration shows a seed, seed head and clonal growth of wood anemones from the rhizome and that clonal growth reveals itself in woodland as communities of wood anemones that flower simultaneously.

Cushion of flowering wood anemones in beech woodland.

I recently chanced upon wood anemone epiphytes on an outsize derelict beech coppice stool base, then even more recently a cushion of wood anemones next to a beech tree and wondered if the dispersal of viable seed was by a woodland bird. With my curiosity duly piqued I consulted Google scholar for research on the seed dispersal mechanisms of wood anemone and discovered that:

1. Wood anemones are a myrmecochorous plant. Ant dispersal of wood anemones are covered in two papers investigating the colonisation of woodlands in Belgium (Bossuyt, B., Hermy, M. and Deckers, J., 1999. Migration of herbaceous plant species across ancient–recent forest ecotones in central Belgium. Journal of Ecology, 87(4), pp.629-638.(4), pp.629-638.) and Sweden (Brunet, J. and Von Oheimb, G., 1998. Migration of vascular plants to secondary woodlands in southern Sweden. Journal of Ecology, 86(3), pp.429-438.  The papers emphasise the slow colonising rates of ancient woodland indicator plants into new woodland.

2. Wood anemone seed is covered in bristles and Belgian researchers established that it can be effectively transported and dispersed in the fur of some animals. Couvreur, M., Verheyen, K. and Hermy, M., 2005. Experimental assessment of plant seed retention times in fur of cattle and horse. Flora-Morphology, Distribution, Functional Ecology of Plants, 200(2), pp.136-147 In my local woodlands badgers (Meles meles), foxes (Vulpes vulpes) squirrels (Sciurus carolinensis), fallow/roe/sika/muntjac deer and wild boar (Sus scrofa) would be likely candidates for the dispersal of wood anemone seeds. Both boar and badgers root around in the soil and it's plausible that a seed could be transported and then deposited in freshly disturbed soil.

3.  A research paper from Germany investigated wood anemone seed dispersal in beech woodland where ants are absent, found that gastropods can ingest wood anemone seeds and then excrete them with minimal adverse impact on the seeds ability to germinate.  Türke, M., Heinze, E., Andreas, K., Svendsen, S.M., Gossner, M.M. and Weisser, W.W., 2010. Seed consumption and dispersal of ant-dispersed plants by slugs. Oecologia, 163(3), pp.681-693.Oecologia, 163(3), pp.681-693. The paper also highlighted the destruction of wood anemone seeds by rodents, effectively ruling out rodents as a vector in seed dispersal.

4. Avian dispersal doesn't appear to feature in any research material. There was also no research on seeds trodden on and then transported on the shoe tread, paw, foot or hoof of whatever had stepped on the seed(s).

So although I am no wiser as to how the wood anemones I photographed were dispersed, I am now aware that wood anemone seeds can be dispersed by ants, gastropods and animals.