Orchid embryo-to-seed (E:S) ratio as an indicator of germination behaviour and its ecological implications

Although the seeds of various Orchidaceae members can be readily germinated on sterile nutrient media, numerous species exhibit complete asymbiotic germination failure. Also, while seed morphology in orchids has been previously linked to dispersal, associations with germinability have not been widely explored. We compiled seed morphometric and germination data for 203 orchid species globally, drawing from international literature and our lab's unpublished findings. Based on (a) final germination percentage and (b) pre-treatment duration, two distinct groups of asymbiotic germination behaviour emerged – termed compliant and defiant, respectively. Additionally, a statistically significant relationship between germinability and the embryo-to-seed (E:S) length ratio was observed. E:S values tended to be lower in species with defiant germination and higher in those with compliant germination. We further correlated seed morphometric and germination data with phylogenetically and ecologically significant parameters (subfamily, growth form, mycoheterotrophy level, climatic zone and habitat shadiness). Notably, lower E:S values and defiant germination predominate in orchids of shaded habitats, while compliant germination is more prevalent in open habitats. Lastly, considering the mycoheterotrophy continuum, we propose that the reduction in the E:S ratio is linked to the adaptation of various orchid lineages to shaded habitats through both an increased dispersal ability and their greater reliance on fungal symbiosis.