Phylogenetic inferences in the dung-moss family Splachnaceae from analyses of cpDNA sequence data and implications for the evolution of entomophily

The moss family Splachnaceae is characterized by half of its members relying on insects for spore dispersal. These species grow on dung or other animal substrates. They produce small and aggregated spores, and their capsule is modified to attract coprophilous insects or carrion flies using olfactory and visual cues. Systematic concepts and implicit evolutionary inferences have relied much on variation in characters associated with the spore dispersal syndrome. Phylogenetic reconstructions based on sequence variation of two chloroplast loci (trnL-trnF region and the rps4 gene) suggest that most supraspecific taxa are poly- or paraphyletic. Transformations in morphological characters associated to the syndrome thus offer little if any phylogenetically informative signal. Brachymitrion is resolved in a nested position within Tayloria. A new combination, Tayloria immersa (Goffinet) Goffinet, Shaw & Cox is proposed for B. immersum. Only one of the five subgenera of Tayloria (subg. Orthodon) is potentially monophyletic. Voitia shares a common ancestor with Tetraplodon and is thus nested within the Splachnoideae. The affinities of Aplodon remain ambiguous. Reconstruction of shifts between wind and insect spore dispersal syndromes suggests that entomophily arose more than once and may have been followed by a reversal to the generalist strategy in two lineages.