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Despite over a century of observations, the obligate insect parasites within the order Entomophthorales remain poorly characterized at the genetic level. In this manuscript, we present a genome for a laboratory-tractableEntomophthora muscaeisolate that infects fruit flies. OurE. muscaeassembly is 1.03 Gb, consists of 7810 contigs and contains 81.3% complete fungal BUSCOs. Using a comparative approach with recent datasets from entomophthoralean fungi, we show that giant genomes are the norm within Entomophthoraceae owing to extensive, but not recent, Ty3 retrotransposon activity. In addition, we find thatE. muscaeand its closest allies possess genes that are likely homologs to the blue-light sensorwhite-collar 1, aNeurospora crassagene that has a well-established role in maintaining circadian rhythms. We uncover evidence thatE. muscaediverged from other entomophthoralean fungi by expansion of existing families, rather than loss of particular domains, and possesses a potentially unique suite of secreted catabolic enzymes, consistent withE. muscae’s species-specific, biotrophic lifestyle. Finally, we offer a head-to-head comparison of morphological and molecular data for species within theE. muscaespecies complex that support the need for taxonomic revision within this group. Altogether, we provide a genetic and molecular foundation that we hope will provide a platform for the continued study of the unique biology of entomophthoralean fungi. 

The millipede Brachycybe lecontii Wood, 1864 is a fungivorous social millipede known for paternal care of eggs and forming multi-generational aggregations. We investigated the life history, paternal care, chemical defence, feeding and social behaviour of B. lecontii and provided morphological and anatomical descriptions, using light and scanning electron microscopy. Based on observations of B. lecontii from 13 locations throughout its distribution, we report the following natural history aspects. The oviposition period of B. lecontii lasted from mid-April to late June and the incubation period lasted 3–4 weeks. Only males cared for the eggs and subsequent care of juveniles was not observed. In one case, the clutches of two males became combined and they were later cared for by only one of the males. The defensive compound of B. lecontii is stored in large glands occupying a third of the paranotal volume and were observed only in stadia II millipedes and older. We observed B. lecontii feeding on fungi of the order Polyporales and describe a cuticular structure on the tip of the labrum that may relate to fungivory. We found that their stellate-shaped aggregations (pinwheels) do not form in the absence of fungus and suggest the aggregation is associated with feeding. We describe and illustrate a previously undescribed comb-like structure on the tibia and tarsi of the six anterior-most leg-pairs and measure the colour and spectral reflectance of the B. lecontii exoskeleton.