Handkea excipuliformis
Encyclopedia
Handkea excipuliformis, synonymous
Synonym (taxonomy)
In scientific nomenclature, a synonym is a scientific name that is or was used for a taxon of organisms that also goes by a different scientific name. For example, Linnaeus was the first to give a scientific name to the Norway spruce, which he called Pinus abies...

 with Lycoperdon polymorphum var. excipuliforme, Calvatia excipuliformis, Calvatia saccata, Lycoperdon cervinum sensu Bolton (1789) p.p., Lycoperdon elatum, Lycoperdon excipuliforme, Lycoperdon excipuliforme forma flavescens, Lycoperdon excipuliforme var. flavescens, Lycoperdon saccatum & Pestle Puffball, is a species of the Lycoperdaceae
Lycoperdaceae
Lycoperdaceae is a family of approximately 150 fungi now known to lie in the Agaricales. Historically they were placed in their own order Lycoperdales. Members of the Lycoperdaceae family are known as the true puffballs. Unlike other types of fungi that hold spores in gills or teeth, puffballs...

 family of puffballs. A rather large mushroom
Mushroom
A mushroom is the fleshy, spore-bearing fruiting body of a fungus, typically produced above ground on soil or on its food source. The standard for the name "mushroom" is the cultivated white button mushroom, Agaricus bisporus; hence the word "mushroom" is most often applied to those fungi that...

, it may reach dimensions of up to 12 cm (4.7 in) broad by 20 cm (7.9 in) tall. It is commonly known
Common name
A common name of a taxon or organism is a name in general use within a community; it is often contrasted with the scientific name for the same organism...

 as the Pestle puffball. or Long-stemmed puffball. Widespread in northern temperate
Temperate
In geography, temperate or tepid latitudes of the globe lie between the tropics and the polar circles. The changes in these regions between summer and winter are generally relatively moderate, rather than extreme hot or cold...

 zones, it is found frequently on pastures and sandy heath
Heath (habitat)
A heath or heathland is a dwarf-shrub habitat found on mainly low quality acidic soils, characterised by open, low growing woody vegetation, often dominated by plants of the Ericaceae. There are some clear differences between heath and moorland...

s.

Taxonomy

This puffball has been variously placed in the genera Bovista
Bovista
Bovista is a genus of fungi in the family Agaricaceae, commonly known as the true puffballs. It was formerly classified within the now-obsolete order Lycoperdales, which, following a restructuring of fungal taxonomy brought about by molecular phylogeny, has been split; the species of Bovista are...

, Lycoperdon
Lycoperdon
Lycoperdon is a genus of puffball mushrooms. The genus has a widespread distribution and contains about 50 species. In general, it contains the smaller species such as the pear shaped puffball and the gem studded puffball. Most of the time they grow from dead wood and if they grow through the...

, Calvatia
Calvatia
Calvatia is a genus of puffball mushrooms which includes the spectacular giant puffball C. gigantea. It was formerly classified within the now-obsolete order Lycoperdales, which, following a restructuring of fungal taxonomy brought about by molecular phylogeny, has been split; the puffballs,...

, and Utraria. In 1989, German mycologist Hanns Kreisel described the genus Handkea to include species of Calvatia that had distinct microscopic features: Handkea
Handkea
Handkea is a genus of puffball mushrooms in the Lycoperdaceae family. In 1989, German mycologist Hanns Kreisel described the genus Handkea to include species of Calvatia with distinct microscopic features, including a unique type of capillitium , with curvy slits instead of the usual pores...

species have a unique type of capillitium (coarse thick-walled hyphae in the gleba
Gleba
Gleba is the fleshy spore-bearing inner mass of fungi such as the puffball or stinkhorn.The gleba is a solid mass of spores, generated within an enclosed area within the sporocarp. The continuous maturity of the sporogenous cells leave the spores behind as a powdery mass that can be easily blown away...

), with curvy slits instead of the usual pores. Although accepted by some authors, the genus concept has been rejected by others.

Phylogenetic analyses published in 2008 shows that Handkea may be grouped in a clade
Clade
A clade is a group consisting of a species and all its descendants. In the terms of biological systematics, a clade is a single "branch" on the "tree of life". The idea that such a "natural group" of organisms should be grouped together and given a taxonomic name is central to biological...

 along with species from several other genera, including Lycoperdon
Lycoperdon
Lycoperdon is a genus of puffball mushrooms. The genus has a widespread distribution and contains about 50 species. In general, it contains the smaller species such as the pear shaped puffball and the gem studded puffball. Most of the time they grow from dead wood and if they grow through the...

, Vascellum, Morganella, Bovistella
Bovistella
Bovistella is a genus of fungi in the family Agaricaceae.-External links:*...

, and Calvatia
Calvatia
Calvatia is a genus of puffball mushrooms which includes the spectacular giant puffball C. gigantea. It was formerly classified within the now-obsolete order Lycoperdales, which, following a restructuring of fungal taxonomy brought about by molecular phylogeny, has been split; the puffballs,...

. Published in the same year, another DNA analysis of the structure of ITS2
Internal transcribed spacer
ITS refers to a piece of non-functional RNA situated between structural ribosomal RNAs on a common precursor transcript. Read from 5' to 3', this polycistronic rRNA precursor transcript contains the 5' external transcribed sequence , 18S rRNA, ITS1, 5.8S rRNA, ITS2, 28S rRNA and finally the 3'ETS...

 rDNA
RDNA
rDNA may stand for:*ribosomal DNA*recombinant DNARDNA is also an acronym for*Reformed Druids of North America...

 transcript
Transcription (genetics)
Transcription is the process of creating a complementary RNA copy of a sequence of DNA. Both RNA and DNA are nucleic acids, which use base pairs of nucleotides as a complementary language that can be converted back and forth from DNA to RNA by the action of the correct enzymes...

 confirmed that H. utriformis is closely related to Lycoperdon echinatum
Lycoperdon echinatum
Lycoperdon echinatum, commonly known as the spiny puffball or the spring puffball, is a type of puffball mushroom in the genus Lycoperdon. The saprobic species has been found in Africa, Europe, Central America and North America, where it grows on soil in deciduous woods, glades, and pastures...

.

Description

Like all puffballs, Handkea excipuliformis has a gasteroid basidiocarp
Basidiocarp
In fungi, a basidiocarp, basidiome or basidioma , is the sporocarp of a basidiomycete, the multicellular structure on which the spore-producing hymenium is borne. Basidiocarps are characteristic of the hymenomycetes; rusts and smuts do not produce such structures...

, meaning the spore
Spore
In biology, a spore is a reproductive structure that is adapted for dispersal and surviving for extended periods of time in unfavorable conditions. Spores form part of the life cycles of many bacteria, plants, algae, fungi and some protozoa. According to scientist Dr...

s are produced internally, and are only released as the mature fruiting body ages and dries, or is broken. Young puffballs are typically 6 to 12 cm (2.4 to 4.7 in) across, white, or pale grey-brown; in maturity it may attain dimensions of 12 centimetres (4.7 in) broad by 20 centimetres (7.9 in) tall. The underside of the puffball is attached to the ground by a root-like assemblage of hyphae called a rhizomorph
Mycelial cord
Mycelial cords are linear aggregations of parallel-oriented hyphae. The mature cords are composed of wide, empty vessel hyphae surrounded by narrower sheathing hyphae...

. This fungus comprises two parts. The upper, globe-like section, which is white at first and turns ochre as it ages, is initially covered in soft, pointed warts; these fall off to leave a smooth, matt surface. Inside this rounded head the spores develop. The brown spores are released into the air; this process is often hastened by rain, or by being trodden on by cattle. Eventually, all that remains is the sterile pestle-shaped base.

The stipe
The stipe expands once the head has ruptured and released the spores and then remain intact throughout the winter and into the following summer. It is parallel or slightly tapering in at the base; spongy; surface soon becoming wrinkled; initially white with pointed warts, but later turning ochre and becoming smooth and leathery.

Spores

Initially the spore mass (gleba) is white, later becoming olive and then purple-brown at maturity. Spores are 3.5 to 5.5 micrometres, warted and ball shaped.

Distribution and habitat

Common and widespread from late Summer until Autumn. The pestle puffball Grows singly or in small groups in humus soil in both coniferous and broadleaf woodland and on short grassland. Frequent beneath hedges, on wasteland and in all kinds of woods; particularly common on the edges of woodland clearings.

Edibility

This fairly large puffball
Puffball
A puffball is a member of any of several groups of fungus in the division Basidiomycota. The puffballs were previously treated as a taxonomic group called the Gasteromycetes or Gasteromycetidae, but they are now known to be a polyphyletic assemblage. The distinguishing feature of all puffballs is...

 is edible only when the spore bearing flesh is young, and white. The taste and odour are not distinctive. Edible only when young and white throughout. It tastes very similar to the Giant Puffball, but the flesh is not quite as firm and the outer skin should be removed.

Similar species

Handkea perlatum
Handkea
Handkea is a genus of puffball mushrooms in the Lycoperdaceae family. In 1989, German mycologist Hanns Kreisel described the genus Handkea to include species of Calvatia with distinct microscopic features, including a unique type of capillitium , with curvy slits instead of the usual pores...

is much smaller, has a shorter stipe, and retains a mesh-like pattern when the warts are rubbed off the cap.
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