Sphaerobolus
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Stages in the liberation of the peridiolum by ''Sphaerobolus stellatus'' ''Sphaerobolus'' is a genus of fungi in the family
Geastraceae Geastrales is an order of gasterocarpic basidiomycetes (fungi) that are related to Cantharellales. The order contains the single family Geastraceae, commonly known as "earthstars", which older classifications had placed in Lycoperdales, or Pha ...
. Commonly known as the "shotgun fungus", "artillery fungus" or "cannonball fungus", species discharge their spores with explosive force. Discharged spore sacs are sticky and have a tendency to strongly adhere to whatever surface they encounter, making them a nuisance to homeowners, pressure washing contractors, landscape
mulch A mulch is a layer of material applied to the surface of soil. Reasons for applying mulch include conservation of soil moisture, improving fertility and health of the soil, reducing weed growth and enhancing the visual appeal of the area. A m ...
producers and insurance companies.


Taxonomy and classification

The generic name is derived from the
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
words ''sphaer'', meaning "sphere", and ''obolus'', meaning "to throw". ''Sphaerobolus'' was first described by the Italian priest and biologist
Pier Antonio Micheli Pier Antonio Micheli (December 11, 1679 – January 1, 1737) was a noted Italian botanist, professor of botany in Pisa, curator of the Orto Botanico di Firenze, author of ''Nova plantarum genera iuxta Tournefortii methodum disposita''. He discover ...
(as ''Carpobolus'') in 1729. Formerly, the genus was placed either in its own family, the Sphaerobolaceae, in the order Sclerodermatales, or, more commonly, in the order Nidulariales. Currently, the genus is placed in the family Geastraceae. Recent
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
analysis suggests that ''Sphaerobolus'' should be placed in the gomphoid-phalloid
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
along with related genera like ''
Geastrum ''Geastrum'' (orthographical variant ''Geaster'') is a genus of puffball-like mushrooms in the family Geastraceae. Many species are known commonly as earthstars. The name, which comes from ''geo'' meaning ''earth'' and meaning ''star'', refers ...
'', ''
Phallus A phallus is a penis (especially when erect), an object that resembles a penis, or a mimetic image of an erect penis. In art history a figure with an erect penis is described as ithyphallic. Any object that symbolically—or, more precisel ...
'', '' Pseudocolus'', ''
Ramaria The genus ''Ramaria'' comprises approximately 200 species of coral fungi. Several, such as '' Ramaria flava'', are edible and picked in Europe, though they are easily confused with several mildly poisonous species capable of causing nausea, vomi ...
'', '' Clavariadelphus'', '' Gomphus'' and '' Gautieria''. Within the genus, three highly supported clades may be discerned, corresponding to ''S. stellatus'', ''S. iowensis'', and the recently described taxon ''S. ingoldii''.


Description

''Sphaerobolus stellatus'', the most common species, is common on wood mulch worldwide.
Basidiocarps In fungi, a basidiocarp, basidiome, or basidioma () is the sporocarp (fungi), sporocarp of a basidiomycota, basidiomycete, the Multicellular organism, multicellular structure on which the spore-producing hymenium is borne. Basidiocarps are chara ...
, which grow in groups, are 1–3 mm in diameter, roughly spherical or ovoid in shape, and white, buff, pale brown or orange in color. The peridium (outer wall) consists of several layers, including a gelatinous layer. At maturity, the exoperidium (outermost layer) splits into several lobes to expose the single, reddish black to dark brown peridiole (spore casing) about 1 mm diam. The peridiole is forcibly ejected (sometimes up to several feet), leaving an evaginated endoperidium.
Spore In biology, a spore is a unit of sexual or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plants, algae, f ...
s develop inside the peridiole and are typically 6–10 x 4–6 
μm The micrometre ( international spelling as used by the International Bureau of Weights and Measures; SI symbol: μm) or micrometer (American spelling), also commonly known as a micron, is a unit of length in the International System of Unit ...
, elliptical to oblong in shape, thick-walled, and
hyaline A hyaline substance is one with a glassy appearance. The word is derived from el, ὑάλινος, translit=hyálinos, lit=transparent, and el, ὕαλος, translit=hýalos, lit=crystal, glass, label=none. Histopathology Hyaline cartilage is ...
, and are intermixed with irregularly-shaped cells called gemmae. Species have a cosmopolitan distribution, and are usually found on dung, decaying wood (such as landscaping mulch), or vegetative litter.


Mechanism of spore discharge

The
peridium The peridium is the protective layer that encloses a mass of spores in fungi. This outer covering is a distinctive feature of gasteroid fungi. Description Depending on the species, the peridium may vary from being paper-thin to thick and rubber ...
, which consists of six distinct layers, forms two "cups" at maturity. Three layers form the outer cup, two form the inner cup, and one layer dissolves to create the fluid that bathes the
gleba Gleba (, from Latin ''glaeba, glēba'', "lump") is the fleshy spore-bearing inner mass of certain fungi such as the puffball or stinkhorn. The gleba is a solid mass of spores, generated within an enclosed area within the sporocarp. The continu ...
. After splitting of the peridal layers to expose the gleba, enzymatic conversion of
glycogen Glycogen is a multibranched polysaccharide of glucose that serves as a form of energy storage in animals, fungi, and bacteria. The polysaccharide structure represents the main storage form of glucose in the body. Glycogen functions as one o ...
to
glucose Glucose is a simple sugar with the molecular formula . Glucose is overall the most abundant monosaccharide, a subcategory of carbohydrates. Glucose is mainly made by plants and most algae during photosynthesis from water and carbon dioxide, using ...
increases the internal
osmotic pressure Osmotic pressure is the minimum pressure which needs to be applied to a solution to prevent the inward flow of its pure solvent across a semipermeable membrane. It is also defined as the measure of the tendency of a solution to take in a pure ...
and the turgidity of palisade cells in the inner peridial cup. Analysis of the glebal
carbohydrates In organic chemistry, a carbohydrate () is a biomolecule consisting of carbon (C), hydrogen (H) and oxygen (O) atoms, usually with a hydrogen–oxygen atom ratio of 2:1 (as in water) and thus with the empirical formula (where ''m'' may or may ...
revealed an increase in
glucose Glucose is a simple sugar with the molecular formula . Glucose is overall the most abundant monosaccharide, a subcategory of carbohydrates. Glucose is mainly made by plants and most algae during photosynthesis from water and carbon dioxide, using ...
,
mannitol Mannitol is a type of sugar alcohol used as a sweetener and medication. It is used as a low calorie sweetener as it is poorly absorbed by the intestines. As a medication, it is used to decrease pressure in the eyes, as in glaucoma, and to lower ...
, and
trehalose Trehalose (from Turkish '' tıgala'' – a sugar derived from insect cocoons + -ose) is a sugar consisting of two molecules of glucose. It is also known as mycose or tremalose. Some bacteria, fungi, plants and invertebrate animals synthesize it ...
prior to glebal discharge, which would account for the increase in osmotic pressure. Glebal discharge typically occurs 5–6 hours after the apex has split. In ''S. stellatus'', the peridiole shoots off with an audible pop, flying up to 6 m, leaving a remnant looking like a miniature earth-star. This species is phototropic, and the nearest source of direct or reflected light will be the target for glebal discharge. The spores remain viable for several years; gemmae can also germinate.


Species

*'' Sphaerobolus ingoldii'' *'' Sphaerobolus iowensis'' *'' Sphaerobolus stellatus'' *'' Sphaerobolus jaysukhianus''


References


External links


Tom Volk's Fungus of the Month, July 2005''Sphaerobolus stellatus''--The Artillery Fungus
* {{Taxonbar, from=Q1393177 Geastraceae