The enigmatic hyphomycete Torula sensu Caspary revisited

About 130 years ago Robert Caspary described fossil fungi resembling the extant anamorphic genus Torula Persoon (Ascomycota, Pezizomycotina) from two pieces of Eocene Baltic amber. Despite being among the earliest fungi recognized from amber, these microfossils have received virtually no attention for the past 100 years. Our recent findings of similar fungal inclusions from Baltic and Bitterfeld amber have revealed that these fungi constituted an abundant, but poorly understood component of these Paleogene amber forests. Here we elucidate the morphology and growth mode of these enigmatic fungi and show that they are clearly distinguished from the moniliform hyphae of capnodialean sooty moulds (Ascomycota, Capnodiales), that they also do not correspond with the extant genus Torula, and cannot with confidence be assigned to any extant genus of dematiaceous hyphomycetes. The life cycle of the fungi involved transitions from vegetative hyphae to conidial states producing non-randomly disarticulating chains of multi-cellular phragmoconidia. We provide an emended description of these fungi and suggest a new fossil genus Casparyotorula that comprises three anamorphic species, Casparyotorula globulifera (Caspary) comb. nov., Casparyotorula heteromorpha (Caspary) comb. nov., and Casparyotorula arnoldii sp. nov.