# Gymnosporangium

Gymnosporangium is a genus of heteroecious plant-pathogenic fungi which alternately infect members of the family Cupressaceae, primarily species in the genus Juniperus (junipers), and members of the family Rosaceae in the subfamily Amygdaloideae (apples, pears, quinces, shadbush, hawthorns, rowans and their relatives). The common name cedar-apple rusts has been used for these fungi. According to the Dictionary of the Fungi (10th edition, 2008), there was 57 species in the genus. In 2023, Species Fungorum lists up to 74 species.
In junipers (the primary hosts, see photo), some species form a ball-like gall about 2–4 cm in diameter which produces a set of orange tentacle-like spore tubes called telial horns. These horns expand and have a jelly like consistency when wet. In other species, such as in G. clarvariforme, the telia are produced directly from the bark of the juniper with no obvious gall formation or swelling. The basidiospores are released and travel on the wind until they infect an apple, pear, hawthorn or suitable tree.
On the secondary hosts, the fungus produces yellowish depressions on the leaves. It also infects the fruit, which grows whitish tubes like a Medusa head. These are the spore tubes. These aeciospores must then infect a juniper to complete the life cycle. All the types of spores can spread over long distances.
The fungus does not cause serious damage to junipers, but apple and pear trees can suffer serious loss of fruit production due to the effects of the fungus. Although the genus has a worldwide distribution, its impact depends on availability of its two host plant species. Individual species are found in Northern and Central America, Asia, Europe and the Middle East.
Due to the economic impacts of the rusts in some areas where orchards are of commercial importance, some regions have attempted to ban the planting of and/or eradicate the coniferous hosts.
In 2020, the monotypic family Gymnosporangiaceae was proposed to accommodate Gymnosporangium. As accepted by Wijayawardene et al. 2020.

## Selected species and hosts

| Species | Primary host | Secondary host | Common name |
| Gymnosporangium amelanchieris | Juniperus sect. Juniperus | Amelanchier |
| Gymnosporangium clavariiforme | Juniperus sect. Juniperus | Amelanchier, Crataegus, Pyrus | Tongues of Fire |
| Gymnosporangium clavipes | Juniperus | Crataegus, Cydonia | Cedar-quince rust, quince rust |
| Gymnosporangium confusum | Juniperus | Crataegus, Cydonia, Mespilus, Pyrus |
| Gymnosporangium cornutum | Juniperus sect. Juniperus | Sorbus subgen. Sorbus | Mountain ash juniper rust |
| Gymnosporangium cupressi | Cupressus | Amelanchier |
| Gymnosporangium dobroznakovii | Juniperus sect. Juniperus | Pyrus |
| Gymnosporangium fusisporum | Juniperus sect. Sabina | Cotoneaster |
| Gymnosporangium gaeumannii | Juniperus communis | (not known) |
| Gymnosporangium globosum | Juniperus | Crataegus | Cedar-hawthorn rust, American hawthorn rust |
| Gymnosporangium gracile | Juniperus | Amelanchier, Crataegus, Cydonia |
| Gymnosporangium harknessianum | Juniperus | Amelanchier |
| Gymnosporangium inconspicuum | Juniperus | Amelanchier |
| Gymnosporangium juniperi-virginianae | Juniperus | Malus | Cedar-apple rust |
| Gymnosporangium kernianum | Juniperus | Amelanchier | Kern's pear rust |
| Gymnosporangium libocedri (Now Gymnotelium blasdaleanum, Pucciniaceae family) | Calocedrus | Amelanchier | Pacific Coast pear rust, Incense cedar broom rust |
| Gymnosporangium malyi | (not known) | Crataegus |
| Gymnosporangium multiporum | Juniperus | (not known) |
| Gymnosporangium nelsonii | Juniperus | Amelanchier | Witches broom rust, Rocky Mountain pear |
| Gymnosporangium nidus-avis | Juniperus sect. Sabina | Crataegus, Cydonia, Malus | Juniper witches' broom rust |
| Gymnosporangium sabinae | Juniperus | Pyrus, Malus, Crataegus | Pear rust, European pear rust, or pear trellis rust |
| Gymnosporangium torminalis-juniperinum | Juniperus sect. Juniperus | Sorbus torminalis |
| Gymnosporangium tremelloides | Juniperus sect. Juniperus | Cydonia, Malus, Sorbus |
| Gymnosporangium yamadae | Juniperus | Malus | Japanese apple rust |

## Full list of species

As accepted by Species Fungorum;
* Gymnosporangium amelanchieris (DC.) E. Fisch. (1909)
* Gymnosporangium annulatum Y.M. Liang & B. Cao (2020)
* Gymnosporangium asiaticum Miyabe ex G. Yamada (1904)
* Gymnosporangium atlanticum Guyot & Malençon (1957)
* Gymnosporangium aurantiacum Syd. & P. Syd. (1904)
* Gymnosporangium bermudianum Earle (1893)
* Gymnosporangium bethelii F. Kern (1907)
* Gymnosporangium biseptatum Ellis (1874)
* Gymnosporangium clavariiforme (Wulfen) DC. (1805)
* Gymnosporangium clavipes Cooke & Peck (1873)
* Gymnosporangium confusum Plowr. (1889)
* Gymnosporangium connersii Parmelee (1965)
* Gymnosporangium corniculans F. Kern (1910)
* Gymnosporangium corniforme Sawada (1928)
* Gymnosporangium cornutum (Pers.) Arthur (1910)
* Gymnosporangium cunninghamianum Barclay (1890)
* Gymnosporangium cupressi Long & Goodd. (1921)
* Gymnosporangium davisii F. Kern (1908)
* Gymnosporangium dobrozrakovae Mitrof. (1969)
* Gymnosporangium echinulatum (Seung K. Lee & Kakish.) Y.M. Liang & B. Cao (2020)
* Gymnosporangium effusum F. Kern (1911)
* Gymnosporangium exiguum F. Kern (1908)
* Gymnosporangium exterum Arthur & F. Kern (1910)
* Gymnosporangium floriforme Thaxt. (1908)
* Gymnosporangium formosanum Hirats. f. & Hashioka (1935)
* Gymnosporangium fraternum F. Kern (1911)
* Gymnosporangium fusisporum E. Fisch. (1918)
* Gymnosporangium gaeumannii H. Zogg (1949)
* Gymnosporangium gjaerumii Korbonsk. & Azbukina (1997)
* Gymnosporangium globosum (Farl.) Farl. (1886)
* Gymnosporangium gracile Pat. (1902)
* Gymnosporangium gracilens (Peck) F. Kern & Bethel (1911)
* Gymnosporangium harknessianum (Ellis & Everh. ex F. Kern) F. Kern (1911)
* Gymnosporangium hemisphaericum Hara (1917)
* Gymnosporangium huanglongense Y.M. Liang & B. Cao (2016)
* Gymnosporangium hyalinum (Cooke) F. Kern (1911)
* Gymnosporangium inconspicuum F. Kern (1907)
* Gymnosporangium juniperi-virginianae Schwein. (1822)
* Gymnosporangium kanasense P. Zhao & L. Cai (2020)
* Gymnosporangium kernianum Bethel (1911)
* Gymnosporangium lachrymiforme P. Zhao & L. Cai (2020)
* Gymnosporangium lianhuaense Y.M. Liang, B. Cao & S.Q. Tao (2020)
* Gymnosporangium meridissimum Crowell (1940)
* Gymnosporangium mespili (DC.) F. Kern (1911)
* Gymnosporangium minus Crowell (1940)
* Gymnosporangium miyabei G. Yamada & I. Miyake (1908)
* Gymnosporangium monticola H.Y. Yun (2009)
* Gymnosporangium multiporum F. Kern (1909)
* Gymnosporangium nelsonii Arthur (1901)
* Gymnosporangium nidus-avis Thaxt. (1891)
* Gymnosporangium niitakayamense Y.M. Shen (2018)
* Gymnosporangium nipponicum G. Yamada (1935)
* Gymnosporangium orientale P. Syd. & Syd. (1912)
* Gymnosporangium padmarense Balf.-Browne (1955)
* Gymnosporangium paraphysatum Vienn.-Bourg. (1961)
* Gymnosporangium pleoporum Y.M. Liang & B. Cao (2020)
* Gymnosporangium przewalskii Y.M. Liang & B. Cao (2017)
* Gymnosporangium raphiolepidis (Syd.) T. Kasuya, K. Hosaka & Kakish. (2020)
* Gymnosporangium sabinae (Dicks.) G. Winter (1884)
* Gymnosporangium shennongjiaense P. Zhao & L. Cai (2020)
* Gymnosporangium shiraianum Hara (1919)
* Gymnosporangium sorbi (Arthur) F. Kern (1911)
* Gymnosporangium spinulosum P. Zhao & L. Cai (2020)
* Gymnosporangium taianum F. Kern (1964)
* Gymnosporangium tiankengense P. Zhao & L. Cai (2020)
* Gymnosporangium tianschanicum Z.Y. Zhao & J.Y. Zhuang (2007)
* Gymnosporangium torminali-juniperini E. Fisch. (1910)
* Gymnosporangium trachysorum F. Kern (1910)
* Gymnosporangium tremelloides R. Hartig (1882)
* Gymnosporangium tsingchenense C.T. Wei (1947)
* Gymnosporangium turkestanicum Tranzschel (1939)
* Gymnosporangium unicorne H.Y. Yun (2009)
* Gymnosporangium vauqueliniae Long & Goodd. (1939)
* Gymnosporangium yamadae Miyabe ex G. Yamada (1904)