Phylogenetic analyses published in 2008 shows that Handkea may be grouped in a clade along with species from several other genera, including Lycoperdon, Vascellum, Morganella, Bovistella, and Calvatia.
[3] Published in the same year, another DNA analysis of the structure of ITS2 rDNA transcript confirmed that H. utriformis is closely related to Lycoperdon echinatum.
[4] Like all puffballs, Lycoperdon excipuliforme has a gasteroid basidiocarp, meaning the spores are produced internally, and are only released as the mature fruiting body ages and dries, or is broken.
The pestle puffball grows singly or in small groups in soil in both coniferous and broadleaf woodland and on short grassland.
[5] The taste and odour are unremarkable, and very similar to the giant puffball, but the flesh is not quite as firm and the outer skin should be removed.