Astraeus hygrometricus, commonly known as the hygroscopic earthstar, the barometer

earthstar, or the false earthstar, is a species of fungus in the family

Diplocystaceae. Young specimens resemble a puffball when young and unopened. In
maturity, the mushroom displays the characteristic earthstar shape that is a result
of the outer layer of fruit body tissue splitting open in a star-like manner. The
false earthstar is an ectomycorrhizal species that grows in association with
various trees, especially in sandy soils. A. hygrometricus has a cosmopolitan
distribution, and is common in temperate and tropical regions. Its common names
refer to the fact that it is hygroscopic (water-absorbing), and can open up its
rays to expose the spore sac in response to increased humidity, and close them up
again in drier conditions. The rays have an irregularly cracked surface, while the
spore case is pale brown and smooth with an irregular slit or tear at the top. The
gleba is white initially, but turns brown and powdery when the spores mature. The
spores are reddish-brown, roughly spherical with minute warts, measuring 7.511
micrometers in diameter.

Despite a similar overall appearance, A. hygrometricus is not related to the true

earthstars of genus Geastrum, although historically, they have been taxonomically
confused. The species was first described by Christiaan Hendrik Persoon in 1801 as
Geastrum hygrometricus. In 1885, Andrew P. Morgan proposed that differences in
microscopic characteristics warranted the creation of a new genus Astraeus distinct
from Geastrum; this opinion was not universally accepted by later authorities.
Several Asian populations formerly thought to be A. hygrometricus were renamed in
the 2000s once phylogenetic analyses revealed they were unique Astraeus species,
including A. asiaticus and A. odoratus. Research has revealed the presence of
several bioactive chemical compounds in the fruit bodies. North American field
guides typically rate A. hygrometricus as inedible.

Contents

1 Taxonomy, naming, and phylogeny

2 Description
2.1 Edibility
3 Similar species
4 Habitat, distribution, and ecology
5 Bioactive compounds
6 Traditional beliefs
7 See also
8 References
9 External link

Taxonomy, naming, and phylogeny

Because this species resembles the earthstar fungi of Geastrum, it was placed in
that genus by early authors, starting with Christian Hendrik Persoon in 1801[2] (as
Geaster, an alternate spelling of Geastrum). According to the American botanist
Andrew P. Morgan, however, the species differed from those of Geastrum in not
having open chambers in the young gleba, having larger and branched capillitium
threads, not having a true hymenium, and having larger spores. Accordingly, Morgan
set Persoon's Geaster hygrometricum as the type species of his new genus Astraeus
in 1889.[3] Despite Morgan's publication, some authorities in the following decades
continued to classify the species in Geastrum.[4][5] The New-Zealand based
mycologist Gordon Herriot Cunningham explicitly transferred the species back to the
genus Geastrum in 1944, explaining:

The treatment of this species by certain taxonomists well illustrates the

pitfalls that lie in wait for those who worship at the shrine of ontogenic
classification ... The only feature of those outlined in which the species differs
from others of Geastrum is the somewhat primitive hymenium. In the developing plant
the glebal cavities are separated by tramal plates so tenuous as to be overlooked
by the uncritical worker. Each cavity is filled with basidia somewhat irregularly
arranged in clusters (like those of Scleroderma) and not in the definite palisade
of the species which have been studied. This difference disappears as maturity is
reached, when plants resemble closely the fructification of any other member of the
genus. The taxonomist is then unable to indicate any point of difference by which
"Astraeus" may be separated from Geastrum, which indicates that the name should be
discarded.[6]

Cunningham's treatment was not followed by later authorities, who largely

considered Astraeus a distinct genus. According to the taxonomical authority
MycoBank,[1] synonyms of Astraeus hygrometricus include Lycoperdon stellatus Scop.
(1772);[7] Geastrum fibrillosum Schwein. (1822);[8] Geastrum stellatum (Scop.)
Wettst. (1885); and Astraeus stellatus E.Fisch. (1900).[9]

Astraeus hygrometricus has been given a number of colloquial names that allude to
its hygroscopic behavior, including the "hygrometer earthstar", the "hygroscopic
earthstar", the "barometer earthstar", and the "water-measure earthstar".[10][11]
The resemblance to Geastrum species (also known as true earthstars) accounts for
the common name "false earthstar".[12] The specific name is derived from the Greek
words ????? (hygros) "wet" and ?t??? (metron) "measure".[13] The German
Mycological Society selected the species as their "Mushroom of the Year" in 2005.
[14]

Studies in the 2000s showed that several species from Asian collection sites
labelled under the specific epithet hygrometricus were actually considerably
variable in a number of macroscopic and microscopic characteristics.[15] Molecular
studies of the DNA sequences of the ITS region of the ribosomal DNA from a number
of Astraeus specimens from around the world have helped to clarify phylogenetic
relationships within the genus. Based on these results, two Asian "hygrometricus"
populations have been described as new species: A. asiaticus[15] and A. odoratus
(synonymous with Petcharat's A. thailandicus described in 2003[16]). Preliminary
DNA analyses suggests that the European A. hygrometricus described by Persoon is a
different species than the North American version described by Morgan, and that the
European population may be divided into two distinct phylotypes, from France and
from the Mediterranean.[17][18] A 2010 study identified a Japanese species,
previously identified as A. hygrometricus, as genetically distinct; it has yet to
be officially named.[18]

A form of the species found in Korea and Japan, A. hygrometricus var. koreanus, was
named by V.J. Stanek in 1958;[19] it was later (1976) published as a distinct
speciesA. koreanusby Hanns Kreisel.[20] As pointed out by Fangfuk and colleagues,
clarification of the proper name for this taxon must await analysis of A.
hygrometricus var. koreanus specimens from the type locality in North Korea.[18]
Description
A collection of fruit bodies

Young specimens of A. hygrometricus have roughly spherical fruit bodies that

typically start their development partially embedded in the substrate. A smooth
whitish mycelial layer covers the fruit body, and may be partially encrusted with
debris. As the fruit body matures, the mycelial layer tears away, and the outer
tissue layer, the exoperidium, breaks open in a star-shaped (stellate) pattern to
form 420 irregular "rays". This simultaneously pushes the fruit body above ground
to reveal a round spore case enclosed in a thin papery endoperidium. The rays open
and close in response to levels of moisture in the environment, opening up in high
humidity, and closing when the air is dry.[21] This is possible because the
exoperidium is made of several different layers of tissue; the innermost, fibrous
layer is hygroscopic, and curls or uncurls the entire ray as it loses or gains
moisture from its surroundings.[22] This adaptation enables the fruit body to
disperse spores at times of optimum moisture, and reduce evaporation during dry
periods.[23][24] Further, dry fruit bodies with the rays curled up may be readily
blown about by the wind, allowing them to scatter spores from the pore as they
roll.[21]
"This veritable barometer is the most theatrical of all the earthstars. A few
minutes immersion in water will open up old, dried-up specimens that seem as
tightly closed as clenched fists."
David Arora in Mushrooms Demystified[25]

The fruit body is 18 cm (0.43.1 in) in diameter from tip to tip when expanded.
[25] The exoperidium is thick, and the rays are typically areolate (divided into
small areas by cracks and crevices) on the upper surface,[26] and are dark grey to
black. The spore case is sessile (lacking a stalk), light gray to tan color and 1
to 3 cm (0.4 to 1.2 in) broad with a felt-like or scurfy (coated with loose scaly
crust) surface; the top of the spore case is opened by an irregular slit, tear or
pore.[27] The interior of the spore case, the gleba, is white and solid when young,
and divided into oval loculesa characteristic that helps to distinguish it from
Geastrum.[28] The gleba becomes brown and powdery as the specimen matures.[29]
Small dark hairlike threads (rhizomorphs) extend from the base of the fruit body
into the substrate. The rhizomorphs are fragile, and often break off after
maturity.[4]
Mature fruit body (left); cross-section of young fruit body (right)

The spores are spherical or nearly so, reddish-brown, thick-walled and verrucose
(covered with warts and spines). The spores' dimensions are 711 m;[30] the warts
are about 1 m long.[31] The spores are non-amyloid, and will not stain with iodine
from Melzer's reagent.[10] The use of scanning electron microscopy has shown that
the spines are 0.901.45 m long, rounded at the tip, narrow, tapered, and sometime
joined together at the top.[15] The capillitia (masses of thread-like sterile
fibers dispersed among the spores) are branched, 3.56.5 m in diameter, and
hyaline (translucent).[4] The basidia (spore-bearing cells) are four- to eight-
spored,[4] with very short sterigmata.[32] The basidia are arranged in long strings
of clusters; individual basidia measure 1115 by 1824 m. The threads of the
capillitia arise from the inner surface of the peridium, and are thick-walled,
long, interwoven, and branched, measuring 35.5 m thick.[4] The exoperidium (the
outer layer of tissue, comprising the rays) is made of four distinct layers of
tissue: the mycelial layer contains branched hyphae that are 46 m in diameter;
the hyphae of the fibrous layer are 68 m diameter and branched; the collenchyma-
type layer has branched hyphae of 34 m diameter; the soft layer contains hyphae
that are 36 m in diameter.[33]
Edibility
Specimen from Japan

North American sources describe A. hygrometricus as being of either unknown

edibility,[26] or too tough to be edible.[27][34][35] However, they are regularly
consumed in Asia, including Nepal[36] and South Bengal, where "local people consume
them as delicious food".[37] They are collected from the wild and sold in the
markets of India.[38][39]

A study of a closely related southeast Asian Astraeus species concluded that the
fungus contained an abundance of volatile eight-carbon compounds (including 1-
octanol, 1-octen-3-ol, and 1-octen-3-one) that imparted a "mushroom-like, earthy,
and pungent odor that was evident as an oily and moss-like smell upon opening the
caps". The study's authors further noted that the fruit bodies after cooking have a
"roasted, maillard, herbal, and oily flavor". Volatile compounds detected after
cooking the mushroom samples included furfural, benzaldehyde, cyclohexenone, and
furanyl compounds.[40] The regional differences in opinions on edibility are from
sources published before it was known that North American and Asian versions of A.
hygrometricus were not always the same; in some cases Asian specimens have been
identified as new species, such as A. asiaticus and A. odoratus.[15][17]
Similar species
Astraeus pteridis is similar in appearance, but larger; it is found in North
America and the Canary Islands.

Although A. hygrometricus bears a superficial resemblance to member of the "true

earthstars" Geastrum, it may be readily differentiated from most by the hygroscopic
nature of its rays. Hygroscopic earthstars include G. arenarium, G. corollinum, G.
floriforme, G. recolligens, and G. kotlabae.[25] Unlike Geastrum, the young fruit
bodies of A. hygrometricus do not have a columella (sterile tissue in the gleba, at
the base of the spore sac).[41] Geastrum tends to have its spore sac opening
surrounded by a peristome or a disc, in contrast with the single lacerate slit of
A. hygrometricus. There are also several microscopic differences: in A.
hygrometricus, the basidia are not arranged in parallel columns, the spores are
larger, and the threads of the capillitia are branched and continuous with the
hyphae of the peridium.[4][26] Despite these differences, older specimens can be
difficult to distinguish from Geastrum in the field.[11] One species of Geastrum,
G. mammosum, does have thick and brittle rays that are moderately hygroscopic, and
could be confused with A. hygrometricus; however, its spores are smaller than A.
hygrometricus, typically about 4 m in diameter.[30]

Astraeus pteridis is larger, 5 to 15 cm (2.0 to 5.9 in) or more when expanded, and
often has a more pronounced areolate pattern on the inner surface of the rays.[25]
It is found in North America and the Canary Islands.[17] A. asiaticus and A.
odoratus are two similar species known from throughout Asia and Southeast Asia,
respectively.[17] A. odoratus is distinguished from A. hygrometricus by a smooth
outer mycelial layer with few adhering soil particles, 39 broad rays, and a fresh
odor similar to moist soil. The spore ornamentation of A. odoratus is also distinct
from A. hygrometricus, with longer and narrower spines that often joined together.
[15] A. asiaticus has an outer peridial surface covered with small granules, and a
gleba that is purplish-chestnut in color, compared to the smooth peridial surface
and brownish gleba of A. hygrometricus. The upper limit of the spore size of A.
asiaticus is larger than that of its more common relative, ranging from 8.7515.2
m.[17] A. koreanus (sometimes named as the variety A. hygrometricus var. koreanus;
see Taxonomy) differs from the more common form in its smaller size, paler fruit
body, and greater number of rays; microscopically, it has smaller spores (between
6.8 and 9 m in diameter), and the spines on the spores differ in length and
morphology.[15] It is known from Korea and Japan.[18]
Habitat, distribution, and ecology

Astraeus hygrometricus is an ectomycorrhizal fungus and grows in association with a

broad range of tree species.[42] The mutualistic association between tree roots and
the mycelium of the fungus helps the trees extract nutrients (particularly
phosphorus) from the earth; in exchange, the fungus receives carbohydrates from
photosynthesis.[37] In North America, associations with oak and pine are usual,[10]
while in India, it has been noted to grow commonly with chir pine (Pinus
roxburghii) and sal (Shorea robusta).[37] The false earthstar is found on the
ground in open fields, often scattered or in groups, especially in nutrient-poor,
sandy or loamy soils.[26][31][32] It has also been reported to grow on rocks,
preferring acid substrates like slate and granite, while avoiding substrates rich
in lime.[43] In Nepal, fruit bodies have been collected at elevations of 3,000 m
(9,800 ft).[44] Fruit bodies typically appear in autumn, although the dry fruit
bodies are persistent and may last up to several years.[30] Gelatinipulvinella
astraeicola is a leotiaceous fungus with minute, gelatinous, pulvinate (cushion-
shaped) apothecia, known to grow only on the inner surface of the rays of dead
Astraeus species, including A. hygrometricus.[45]

The species has a cosmopolitan distribution except for arctic, alpine and cold
temperate regions;[32] it is common in temperate and tropical regions of the world.
[17] It has been collected in Africa, Asia, Australia, Europe, North America,[16]
and South America.[46]
Bioactive compounds
Backbone chemical structure upon which the unique triterpenes astrahygrol (R1=OH,
R2=H), 3-epi-astrahygrol (R1=H, R2=OH), and astrahygrone (R1=R2=O) are formed

Mushroom polysaccharides from a number of species have attracted research interest

for their immunomodulatory and antitumor properties.[47] Extracts from A.
hygrometricus containing the polysaccharide named AE2 were found to inhibit the
growth of several tumor cell lines in laboratory tests,[48][49] and stimulated the
growth of splenocytes, thymocytes, and bone marrow cells from mice. The extract
also stimulated mouse cells associated with the immune system; specifically, it
enhanced the activity of mouse natural killer cells, stimulated macrophages to
produce nitric oxide, and enhanced production of cytokines.[38][50][51][52] The
activation of macrophages by AE2 might be mediated by a mitogen-activated protein
kinase pathway of signal transduction.[53][54] AE2 is made of the simple sugars
mannose, glucose, and fucose in a 1:2:1 ratio.[37]

In addition to the previously known steroid compounds ergosta-7,22-diene-3-ol

acetate and ergosta-4,6,8-(14),22-tetraene-3-one, three unique
triterpenesderivatives of 3-hydroxy-lanostanehave been isolated from fruit bodies
of A. hygrometricus. The compounds, named astrahygrol, 3-epi-astrahygrol, and
astrahygrone (3-oxo-25S-lanost-8-eno-26,22-lactone), have d-lactone (a six-membered
ring) in the side chaina chemical feature previously unknown in the
Basidiomycetes.[55][56] A previously unknown steryl ester (3, 5a-dihydroxy-(22E,
24R)-ergosta-7,22-dien-6a-yl palmitate) has been isolated from mycelia grown in
liquid culture. The compound has a polyhydroxylated ergostane-type nucleus.[57]

Ethanol extracts of the fruit body are high in antioxidant activity, and have been
shown in laboratory tests to have anti-inflammatory activity comparable to the drug
diclofenac.[58] Studies with mouse models have also demonstrated hepatoprotective
(liver-protecting) ability, possibly by restoring diminished levels of the
antioxidant enzymes superoxide dismutase and catalase caused by experimental
exposure to the liver-damaging chemical carbon tetrachloride.[59]
Traditional beliefs

This earthstar has been used in traditional Chinese medicine as a hemostatic agent;
the spore dust is applied externally to stop wound bleeding and reduce chilblains.
[60] Two Indian forest tribes, the Baiga and the Bharia of Madhya Pradesh, have
been reported to use the fruit bodies medicinally. The spore mass is blended with
mustard seed oil, and used as a salve against burns.[61] The Blackfoot of North
America called the fungus "fallen stars", considering them to be stars fallen to
the earth during supernatural events.[62]
See also

Fungi portal

Medicinal mushrooms

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External link
Wikimedia Commons has media related to Astraeus hygrometricus.
Wikispecies has information related to: Astraeus hygrometricus

Astraeus hygrometricus in Index Fungorum

Taxon identifiers

Wd: Q980655 BioLib: 60773 EoL: 1029271 GBIF: 5239766 iNaturalist: 55271 Index
Fungorum: 122650 ITIS: 14099 MycoBank: 122650 NCBI: 150782

Categories:

BoletalesEdible fungiFungi described in 1801Fungi of AsiaFungi of

AustraliaFungi of EuropeFungi of North AmericaFungi of South AmericaMedicinal
fungiTaxa named by Christiaan Hendrik Persoon

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