977:: The famous "death grip" exhibited by the ant is also a result of fungus-induced manipulation. This behavior consists of an infected ant locking its mandibles onto (i.e. biting) a leaf so tightly that the ant is prevented from falling as it dies hanging upside down, consequently enabling the proper growth of the fungus' fruiting body. This is possibly a result of the atrophy of the ant's mandibular muscles caused by the secretion of fungal compounds. In multiple studies, fungal cell populations were found within atrophied mandibular muscle tissues. These fungal cell populations have been found to be interconnected through anastomosing tubules. Researchers have proposed that these hyperconnected fungal networks may indicate cooperativity between fungal cells to control the actions of the ant's mandibular muscle. Another common hypothesis amongst researchers is that fungal cells infiltrate between the muscle fibers and then secrete chemicals which cause the muscles to atrophy. Significant decreases in
652:
882:
44:
899:. The behavioral manipulation of the ant, which gives rise to the name "zombie-ant", is an extended phenotype of the fungus. It first affects the ant's behavior through convulsions that make it fall from its high canopy nest onto the forest floor. This is followed by the fungus controlling the climbing of the ant and the locking of its jaw (and subsequent death) onto a leaf around 25 centimetres above the ground, which is thought to be the optimal height for fungal spore growth and dispersion.
65:
1269:
followed by a "death grip" of the infected ant once it has reached a location with optimal conditions for post-mortem fungal development. This leads to the fungus continuing its growth and releasing fungal spores onto the forest floor. These spores will then be encountered by the ants which, when the aerial foraging route is not possible, have to occasionally descend to ground level. Therefore,
854:. The ant is no longer able to control the muscles of the mandible and remains fixed in place, hanging upside-down on the leaf. This lockjaw trait is popularly known as the death grip and is essential in the fungus's lifecycle. A study led in Thailand revealed that there is a synchronization of this manipulated biting behavior at solar noon.
1066:-infected ants die, they are mainly located in regions containing a high density of ants which were previously manipulated and killed. These areas are termed "graveyards" and can be of 20 to 30 metres (66 to 98 ft) in range, with a local density of dead ants possibly exceeding 25 per square metre (2/sq ft).
1355:
three to five meters before going back up into the canopy. This demonstrates the avoidance of the zones of infection by the ants. Additionally, more evidence participates in the favour of this defence method being adaptive as it is not observed in undisturbed forests where the zombie-ant fungus is not present.
1268:
typically depicts the infected ant leaving its canopy nest and its normal foraging path to reach the forest floor and subsequently climbs to around 25 centimetres (9.8 in) above ground level, a height that is considered optimal for fungal growth due to its humidity level and temperature. This is
1256:
Some parasites have evolved to manipulate their host's behavior in order to increase their transmission to uninfected susceptible individuals, thereby increasing their fitness. This host manipulation is termed the "extended phenotype" of the parasite and is a form of adaptation. Host ant manipulation
981:
concentration and mitochondria number were identified in infected ants. A deficit in leucine results in the prevention of muscle regeneration because the amino acid is a nutrient regulator of muscle protein synthesis. A decrease in mitochondria ultimately results in a reduction of energy and calcium
964:
and guanidinobutyric acid (GBA), have been identified as responsible for the manipulation of the host brain. Both compounds are known to be involved in various neurological disorders. However, more research is needed to determine whether other fungal metabolites interact with the host brain to cause
902:
Throughout the lifecycle, unique challenges must be met by equally unique metabolic activities. The fungal pathogen must attach securely to the arthropod exoskeleton and penetrate it—avoiding or suppressing host defenses—then, control the behavior of the host before killing it; and finally, it must
1354:
builds its nests high in the canopy, and has a broad network of aerial trails. These trails occasionally move down to the ground level, where infection and graveyards occur, due to canopy gaps too difficult for the ants to cross. When the trails descend to the forest floor, their length is only of
906:
The behavioral manipulation of the ant would not be possible without the presence of huge fungal cell populations beside the host's brain and within muscles because these lead to the secretion of various metabolites known to have important behavioral consequences. During the infection the parasite
1076:
effects on the host population. In fact, studies have described seasonal patterns in the density of previously infected dead ants, with an increase during the rainy season and a decrease during the dry season. It is thought that large precipitation events at the beginning and the end of the rainy
1015:
on the host have been found to vary according to host species. The ant species which are normally found infected in nature exhibit a manipulated behavior, whereas the species which are not typically infected are killed by the infection, but their behavior is not altered. This is likely due to the
1415:
is similarly able to infect the human population through exchange of bodily fluids, leading to an apocalyptic world inhabited by zombie-like "hungries" who attack non-infected. The novel “The Genius Plague” by David Walton, though not about this species specifically, expands on the idea of fungi
476:
and germination). Moreover, other traits such as the host and the location of the death grip were added to the analyses. The morphological study led to 15 new identified species, with 14 which were distributed in the core clade, and one in the subclade. Moreover, it was found that species in the
1238:
is very virulent, only about 6.5% of all fruiting bodies are viable spore producers. This is caused by the weakening of the fungus by the hyperparasite, which may limit the viability of infectious spores. Ants also groom each other to combat microscopic organisms that could potentially harm the
778:
asexual morph. As for the core clade, these species are also recognizable through the hosts they infect, which are usually neotropical ant species. The subclade does not present the same extended phenotype with the famous "death grip" that O. unilateralis species typically exhibit. Their hosts
718:
species exhibit morphological variations which are most certainly due to their wide geographic range, from Japan to the
Americas. Moreover, it has been hypothesized that their morphological variations may also be a result of one fungus species maximizing its infection on one specific host ant
826:
cm above the forest floor, on the northern side of the plant, in an environment with 94–95% humidity and temperatures between 20 and 30 °C (68 and 86 °F). Infections may lead to 20 to 30 dead ants per square meter. When the dead ants are moved to other places and positions, further
821:
The changes in the behavior of the infected ants are very specific, giving rise to the popular term "zombie ants." Behaviors are tuned for the benefit of the fungus in terms of its growth and its transmission, thereby increasing its fitness. The ant climbs up the stem of a plant and uses its
915:
reacts heterogeneously by secreting different metabolites according to the host tissue it encounters and whether they are live or dead. The identification of these natural products is important in order to understand which aspects of the ants are under control and consequently how
1429:. A mushroom grows out of his head, causing him to act erratically and obsess over hallucinogenic mushrooms. Upon returning to his lair after completing his quest line, the player finds him dead on the ground, with the fungus on his head split in two to spread its spores.
971:, present at high extracellular concentrations. Hypoxanthine has deleterious effects on neural tissues of the cerebral cortex, which in the context of zombie ants may indicate a way for the fungus to alter the motor neurons of the ant, consequently affecting its behavior.
1163:
has been studied as a dye for food, cosmetic, and pharmaceutical manufacturing processes. In fact, naphthoquinone derivatives produced by the fungus show a red color under acidic conditions, and a purple color under basic conditions. These pigments are stable against
281:
for, among other reasons, the production of substances active as antibacterial agents that protect the fungus-host ecosystem against further pathogenesis during fungal reproduction. Because of this secondary metabolism, an interest in the species has been taken by
257:
being characterized by alteration of the behavioral patterns of the infected ant. Infected hosts leave their canopy nests and foraging trails for the forest floor, an area with a temperature and humidity suitable for fungal growth; they then use their
1334:
principal hosts evolved efficient behavioral forms of social immunity. The ants clean each other's exoskeletons to decrease the presence of spores attached. Also, ants can sense that a member of the colony is infected; healthy ants carry the
1312:
fortifies the ant cadaver to prevent its decay, which consequently ensures the growth of the fruiting body. Therefore, the zombie-ant fungus adapts to the short viability of its spores by increasing their production using the dead ant.
1531:
Mongkolsamrit S, Kobmoo N, Tasanathai K, Khonsanit A, Noisripoom W, Srikitikulchai P, et al. (November 2012). "Life cycle, host range and temporal variation of
Ophiocordyceps unilateralis/Hirsutella formicarum on Formicine ants".
869:
grow from the ant's head and rupture, releasing the spores. This process takes 4–10 days. Dead ants are found in areas termed "graveyards" which contain high densities of dead ants previously infected by the same fungus.
750:
asexual morph, which arises from the dorsal neck region of the dead ant and produces a dark brown perithecia attached to its stalk. These species are also recognizable through the host species they infect, which are only
1247:
In host–parasite dynamics, both the host and the parasite are under selective pressure: the parasite evolves to increase its transmission, whereas the host evolves to avoid and/or resist the infection by the parasite.
467:
Further analyses were conducted using a set of different traits. Morphological traits were used and included both macro-morphological characters (e.g. typical single stroma arising from the host's dorsal pronotum, the
447:
species were described based on classic taxonomic criteria, and macro-morphological data with a deeper focus on ascospore and asexual morphology. The asexual morphologies made it possible to distinguish two different
2743:
Amnuaykanjanasin A, Panchanawaporn S, Chutrakul C, Tanticharoen M (August 2011). "Genes differentially expressed under naphthoquinone-producing conditions in the entomopathogenic fungus
Ophiocordyceps unilateralis".
1339:-infected individual far away from the colony to avoid exposure to spores. There are also reports that most worker ants remain inside the nest boundaries; this would mean only foragers were at risk of infection.
262:
to attach themselves to a major vein on the underside of a leaf, where the host remains after its eventual death. The process, leading up to mortality, takes 4–10 days, and includes a reproductive stage where
423:
Support for this term has become increasingly important. In 2011, it was hypothesized that the zombie-ant fungus could actually be described as a complex of species which are host-specific, meaning that one
791:
lives in the high canopy and has an extensive network of aerial trails. Sometimes the canopy gaps are too difficult to cross, so the ants' trails descend to the forest floor where they are exposed to
936:
has to overcome to have a successful infection is to attach itself onto the ant's cuticle and then infiltrate it. For this purpose, the fungus' hypha pierces the exoskeleton using enzymes such as
954:: After the fungus enters the ant, it propagates, and fungal cells are found beside the host's brain. Once the population is of sufficient size, the fungus secretes compounds and takes over the
676:
region of the ant once it is dead. Moreover, perithecia, the spore-bearing sexual structure, can be observed on the stalk, just below its tip. This complex forms the fungus' fruiting body.
1003:
More in-depth research is needed for the identification of other fungal compounds which act to atrophy the mandibular muscles, and for the understanding of their exact effects on the ant.
770:
subclade, as described in 2018, also has distinct morphological characteristics. Its species produce a stroma that grows laterally from the host's thorax which itself generates an orange
2779:
Unagul P, Wongsa P, Kittakoop P, Intamas S, Srikitikulchai P, Tanticharoen M (April 2005). "Production of red pigments by the insect pathogenic fungus
Cordyceps unilateralis BCC 1869".
818:
to manipulate the behavioral patterns exhibited by the ant. An infected ant exhibits irregularly timed full-body convulsions that dislodge it from its canopy nest to the forest floor.
1096:, as well as several structurally uncharacterised substances. These natural products are reportedly being investigated as potential leads in discovery efforts toward immunomodulatory,
366:. However, in 2007, important new molecular data was tested, and enabled them to reorganize the family Clavicipitaceae. It was found that Clavicipitaceae was in fact three distinct
1387:, takes control of humans (as opposed to insects) as an alternative host and causes them to exhibit erratic behaviors, such as the desire to attack and infect non-infected humans.
1069:
The density of dead ants within these graveyards can vary according to climatic conditions. This means that environmental conditions such as humidity and temperature can influence
1383:
is revealed to be the primary cause of the infected outbreak and subsequent collapse of human civilization. In the show, the fungus, having adapted to higher temperatures due to
271:
is, in turn, also susceptible to fungal infection itself, an occurrence that can limit its impact on ant populations, which has otherwise been known to devastate ant colonies.
1146:-methyl erythrostominone, epierythrostominol, deoxyerythrostominol, and 3,5,8-trihydroxy-6-methoxy-2-(5-oxohexa-1,3-dienyl)-1,4-naphthoquinone, which have shown activity in
3052:
1176:, which makes them applicable for food coloring and as a dye for other materials. These attributes also make it a prime candidate for antituberculosis testing in secondary
865:
then sprout out of the ant, securely anchoring it to the plant substrate while secreting antimicrobials to ward off competition. When the fungus is ready to reproduce, its
1308:
investment (growth/survival) by the parasite in order to sustain the growth of the fungus' fruiting body on its host, thereby enabling successive reproduction. To do so,
2647:
1156:
discovery. In addition to having antimalarial activities, all six of these secondary metabolites have been demonstrated to have anticancer and antibacterial activities.
668:
The zombie-ant fungus is easily identifiable when its reproductive structure becomes apparent on its dead host, usually a carpenter ant. At the end of its life cycle,
2404:"From behavior to mechanisms: an integrative approach to the manipulation by a parasitic fungus (Ophiocordyceps unilateralis s.l.) of its host ants (Camponotus spp.)"
436:. Following this, a study conducted in Brazil delimited, using morphological comparisons of the ascospores, germination processes, and asexual morphs, four different
1204:, and antifungal treatments. It has also been reported that polyketides have other therapeutic effects such as antitumor, antioxidant and antiaging activities.
822:
mandibles with abnormal force to secure itself to a leaf vein, leaving dumbbell-shaped marks on it. The ants generally clamp to a leaf's vein at a height of 26
1325:
evolved adaptive behaviors able to limit the contact rate between uninfected susceptible hosts and infected hosts, thereby reducing the risk of transmission.
2708:
Wongsa P, Tasanatai K, Watts P, Hywel-Jones N (August 2005). "Isolation and in vitro cultivation of the insect pathogenic fungus
Cordyceps unilateralis".
1917:
1700:"Hidden diversity behind the zombie-ant fungus Ophiocordyceps unilateralis: four new species described from carpenter ants in Minas Gerais, Brazil"
807:; despite this, the fungus may parasitize other closely related species of ants with lesser degrees of host manipulation and reproductive success.
3027:
1297:
species as they can produce and release within the air, clear and thin-walled spores which are susceptible to environmental conditions such as
873:
The term "zombie ants" has been used in popular media as well as scientific articles, but has also been described as "catchy, yet misleading."
2677:
Kittakoopa P, Punyaa J, Kongsaeree P, Lertwerawat Y, Jintasirikul A, Tanticharoena M, Thebtaranonth Y (1999). "Bioactive naphthoquinones from
3412:
2882:"Insect-specific polyketide synthases (PKSs), potential PKS-nonribosomal peptide synthetase hybrids, and novel PKS clades in tropical fungi"
1077:
season stimulates fungal development, which leads to more spores being released and ultimately more individuals being infected and killed.
1046:
A 48-million-year-old fossil of a leaf stem exhibiting dumbbell-shaped marks characteristic of those made by an ant in the death-grip of
428:
species can only successfully infect and manipulate one host ant species. There is a possibility that this resulted in or reinforced the
2822:
Isaka M, Kittakoop P, Kirtikara K, Hywel-Jones NL, Thebtaranonth Y (October 2005). "Bioactive substances from insect pathogenic fungi".
1119:
replaces the symbiotic bacteria within the cicadas to help the host process sap as nutrients, unlike other related species, such as the
2309:
1830:"Ophiocordyceps unilateralis: A keystone species for unraveling ecosystem functioning and biodiversity of fungi in tropical forests?"
3164:"Three-dimensional visualization and a deep-learning model reveal complex fungal parasite networks in behaviorally manipulated ants"
2859:"A Retrospective Analysis of Cordyceps Anti-Tuberculosis Capsule Combined with Chemotherapy for 614 Cases of Secondary Tuberculosis"
3360:
1373:
has evolved to infect humans, thus creating zombie-like enemies in the game. Also, in episode two of the 2023 television series
651:
755:
species. Once the host is killed by the fungus, it is commonly found fixed through their mandibles onto the surfaces of leaves.
3076:"Gene expression during zombie ant biting behavior reflects the complexity underlying fungal parasitic behavioral manipulation"
1391:, who co-wrote and produced the series, said that everything the series suggests fungi do, they have done forever in real life.
2655:
3302:
3226:
1469:
472:(perithecia) growing from the stroma) and microscopic traits (e.g. the morphology of the ascospores in terms of size, shape,
3552:
827:
vegetative growth and sporulation either fails to occur or results in undersized and abnormal reproductive structures. In
1425:
features an ant character named Sozo who is implied to be under the influence of a parasitic fungus similar in nature to
799:
and eventually break through using mechanical pressure and enzymes. Like other fungi pathogenic to insects in the genus
335:. There have been many debates about whether the zombie-ant fungus (and other fungi) belonged to one or to the other as
1400:
2344:"Specialist and Generalist Fungal Parasites Induce Distinct Biochemical Changes in the Mandible Muscles of Their Host"
354:. The classification was based on different morphological characteristics such as filiform ascospores and cylindrical
17:
691:). These are different in terms of their function and characteristics. Generally, the asexual morphs identified for
3542:
3517:
3417:
440:
species. Afterwards, three new species were described in the
Brazilian Amazon, six in Thailand, and one in Japan.
3532:
746:
core clade, as described in 2018, has distinct morphological characteristics. It exhibits a single stroma with a
2611:
Xiao JH, Zhong JJ (2007). "Secondary metabolites from
Cordyceps species and their antitumor activity studies".
1416:
influencing animals with a fungus that invades human brains and influences their actions toward its advantage.
1346:, provided evidence for the avoidance of the forest floor by the host ants as a defence method. In areas where
1885:
3537:
1293:
has another possible form of adaptation which ensures its repeated reproduction. This would be crucial for
1234:
The graveyards of dead ants are numerous and spread throughout the surrounding area of the colony. Though
810:
Yeast stages of the fungus spread in the ant's body and presumably produce compounds that affect the ant's
64:
723:
of ant can occur within the same area, which means that in order to coexist they have to occupy different
3326:
3321:
3121:"Genetic Underpinnings of Host Manipulation by Ophiocordyceps as Revealed by Comparative Transcriptomics"
1395:
259:
1134:
derivatives are an example of secondary metabolite with important pharmaceutical potentials produced by
3074:
de Bekker C, Ohm RA, Loreto RG, Sebastian A, Albert I, Merrow M, Brachmann A, Hughes DP (August 2015).
1375:
383:, into Ophiocordycipitaceae. Fungi able to parasitize ants were also included in the transfer, such as
1942:
1916:
Andersen SB, Gerritsma S, Yusah KM, Mayntz D, Hywel-Jones NL, Billen J, et al. (September 2009).
895:
life cycle includes and depends on the infection and the manipulation of a carpenter ant, principally
831:, the typical behavior of zombie ants is to attach themselves to the lower side of twigs, not leaves.
3481:
3352:
1490:
1231:
as the fungal stalk emerges from the ant's body, which can stop the stalk from releasing its spores.
958:(CNS), which enables it to manipulate the ant to reach the forest floor and climb up the vegetation.
391:. Following this study, multiple traits such as the production of darkly pigmented, hard to flexible
672:
typically generates a single, wiry yet pliant, darkly pigmented stroma which arises from the dorsal
3547:
3522:
3404:
1036:
ecosystems. However, there are some reports of the zombie-ant fungus in warm-temperate ecosystems.
1016:
heterogeneous nature of the fungus which secretes different metabolites according to host species.
885:
Schematic representation of the ant behavioral manipulation caused by natural products secreted by
881:
3562:
2138:
2087:
de Bekker C, Quevillon LE, Smith PB, Fleming KR, Ghosh D, Patterson AD, Hughes DP (August 2014).
230:
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controls the ant's behavior and this manipulation represents an adaptation for the fungus where
3527:
3264:
3162:
Fredericksen MA, Zhang Y, Hazen ML, Loreto RG, Mangold CA, Chen DZ, Hughes DP (November 2017).
2880:
Amnuaykanjanasin A, Phonghanpot S, Sengpanich N, Cheevadhanarak S, Tanticharoen M (June 2009).
2858:
2554:"Graveyards on the move: the spatio-temporal distribution of dead ophiocordyceps-infected ants"
1181:
983:
955:
851:
429:
3053:"'A growing threat to human health': we are ill-equipped for the dangers of fungal infections"
1459:
1039:
Its distribution includes tropical rainforests located in Brazil, Australia and
Thailand, and
3440:
1996:"Behavioral mechanisms and morphological symptoms of zombie ants dying from fungal infection"
1196:
also produces polyketides. These secondary metabolites have been used in antibiotics such as
1123:, which is a traditional immune booster and cancer treatment in Tibetan and Chinese culture.
1093:
838:, which is 48 million years old. Once the mandibles of the ant are secured to the leaf vein,
234:
159:
3430:
1219:, reported in the lay press as the "antizombie-fungus fungus", that results in only 6–7% of
779:
usually die at the base of large trees in the
Amazonian rainforest, among the moss carpets.
727:. Consequently, the fungi may have evolved at the subspecies level in order to maximize its
373:
The new molecular phylogenetics studies contradicted the older classification and moved all
176:
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2980:
2893:
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In fact, studies suggest that the short viability of the fungal spores lead to the need of
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8:
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1995:
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728:
724:
264:
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1994:
Hughes DP, Andersen SB, Hywel-Jones NL, Himaman W, Billen J, Boomsma JJ (May 2011).
1967:
834:
A search of plant-fossil databases revealed similar marks on a fossil leaf from the
3339:
3219:
Entangled Life: How Fungi Make Our Worlds, Change Our Minds & Shape Our
Futures
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2007:
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Sung GH, Hywel-Jones NL, Sung JM, Luangsa-Ard JJ, Shrestha B, Spatafora JW (2007).
1541:
1420:
1040:
828:
291:
2552:
Pontoppidan MB, Himaman W, Hywel-Jones NL, Boomsma JJ, Hughes DP (12 March 2009).
3467:
2993:
2967:
Andersen SB, Ferrari M, Evans HC, Elliot SL, Boomsma JJ, Hughes DP (2 May 2012).
2578:
1724:
1662:
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908:
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413:
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families: the Clavicipitaceae, the Cordycipitaceae and the Ophiocordycipitaceae.
347:
283:
238:
106:
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3347:
3168:
Proceedings of the National Academy of Sciences of the United States of America
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331:
136:
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2792:
2721:
2105:
2046:
1951:
1545:
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invade more soft tissues and structurally fortify the ant's exoskeleton. More
412:, which means 'in the broad sense', because the species actually represents a
320:
Throughout history there has been confusion about the distinction between the
43:
3557:
3511:
3287:
2142:
2054:
1502:
1216:
379:
286:
chemists, with corresponding discovery of small molecule agents (e.g. of the
3188:
2342:
Zheng S, Loreto R, Smith P, Patterson A, Hughes D, Wang L (September 2019).
2012:
1787:
907:
comes across an array of environments such as different host tissues or the
3207:
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3111:
3012:
2923:
2843:
2800:
2765:
2729:
2632:
2597:
2527:
2509:
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2429:
2379:
2295:
2277:
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2180:
2124:
2031:
1959:
1890:
1863:
1805:
1743:
1681:
1607:
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1553:
1305:
1298:
1177:
1101:
968:
847:
734:
703:
367:
254:
3313:
3243:
3136:
2089:"Species-specific ant brain manipulation by a specialized fungal parasite"
1918:"The life of a dead ant: the expression of an adaptive extended phenotype"
1239:
colony. Additional fungi also grant beneficial assistance to the colony.
1180:
patients, by improving symptoms and enhancing immunity when combined with
758:
3399:
3373:
3296:
2905:
2420:
2403:
2360:
1574:"Phylogenetic classification of Cordyceps and the clavicipitaceous fungi"
1388:
1088:
are known in the pharmaceutical world to be a medically-important group.
1029:
961:
796:
482:
351:
250:
116:
2492:
Hughes, David P.; Wappler, Torsten; Labandeira, Conrad C. (2011-02-23).
2262:"When fiction becomes fact: exaggerating host manipulation by parasites"
3365:
2460:
1220:
1051:
835:
720:
697:
684:
433:
287:
96:
3378:
3119:
Will I, Das B, Trinh T, Brachmann A, Ohm RA, de Bekker C (July 2020).
3028:"Could the Zombie Fungus in TV's The Last of Us Really Infect People?"
2835:
1845:
1530:
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1019:
937:
843:
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325:
180:
76:
3258:
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395:
were defined as characteristics of the family Ophiocordycipitaceae.
362:
were first classified, there was no concrete evidence for the genus
267:
grow from the ant's head, rupturing to release the fungus's spores.
3489:
3461:
3386:
3281:
1933:
1464:
1169:
1148:
945:
811:
688:
673:
2821:
1261:
represents one of the best-known examples of extended phenotypes.
346:
comprises over 400 species, historically classified in the family
3247:
2445:"Zombie ant death grip due to hypercontracted mandibular muscles"
1491:"After This Fungus Turns Ants Into Zombies, Their Bodies Explode"
1197:
987:
978:
862:
839:
1993:
2707:
1227:
inflicts on ant colonies. The hyperparasite moves in to attack
1138:. Six known naphthoquinone derivatives have been isolated from
995:
941:
858:
771:
469:
449:
86:
2086:
1764:"Epitypification and re-description of the zombie-ant fungus,
1571:
2969:"Disease dynamics in a specialized parasite of ant societies"
1278:
1159:
Moreover, the use of red naphthoquinone pigments produced by
991:
355:
321:
3161:
2778:
2494:"Ancient death-grip leaf scars reveal ant–fungal parasitism"
2165:"Ancient death-grip leaf scars reveal ant-fungal parasitism"
1915:
1886:"'Zombie ants' controlled by parasitic fungus for 48m years"
3073:
1165:
857:
The fungus then kills the ant and continues to grow as its
2966:
2938:"The Zombie-Ant Fungus Is Under Attack, Research Reveals"
2341:
903:
protect the carcass from microbial and scavenger attack.
398:
246:
1762:
Evans HC, AraĂşjo JP, Halfeld VR, Hughes DP (June 2018).
1761:
1646:"Ophiocordyceps. I. Myrmecophilous hirsutelloid species"
3253:
2491:
2162:
2139:"Fossil Reveals 48-Million-Year History of Zombie Ants"
1643:
3254:
An Electronic Monograph of Cordyceps and Related Fungi
2781:
Journal of Industrial Microbiology & Biotechnology
2163:
Hughes DP, Wappler T, Labandeira CC (February 2011) .
1644:
AraĂşjo JP, Evans HC, Kepler R, Hughes DP (June 2018).
485:
ants, whereas species in the core clade specialise on
3118:
2401:
1006:
405:
The fungus's scientific name is sometimes written as
277:
and related species are known to engage in an active
2873:
1142:, namely erythrostominone, deoxyerythrostominone, 4-
1526:
1524:
1522:
1520:
1518:
2648:"How a parasitic fungus turns ants into 'zombies'"
2547:
2545:
2543:
2541:
2539:
2537:
1020:Geographic distribution and first known appearance
846:connections in the muscle fibers and reducing the
1828:Evans HC, Elliot SL, Hughes DP (September 2011).
1827:
1697:
774:. Moreover, species within this subclade share a
3509:
2962:
2960:
2958:
2956:
2954:
2082:
2080:
2078:
2076:
2074:
2072:
2070:
1515:
2534:
2397:
2395:
2393:
2391:
2389:
2337:
2335:
2333:
2331:
2329:
290:family) of potential interest for use as human
237:in 1859, and currently found predominantly in
2856:
2402:de Bekker C, Merrow M, Hughes DP (July 2014).
2210:
2208:
1989:
1987:
1985:
1983:
1981:
1979:
1977:
1342:Moreover, one of the fungus' principal hosts,
803:, the fungus targets a specific host species,
719:species (host-specific infections). Different
309:
2951:
2067:
1823:
1821:
1819:
1817:
1815:
1698:Evans HC, Elliot SL, Hughes DP (March 2011).
1693:
1691:
1639:
1637:
1126:
707:, two genera of asexually reproducing fungi.
2815:
2386:
2326:
2044:
1635:
1633:
1631:
1629:
1627:
1625:
1623:
1621:
1619:
1617:
1043:found in South Carolina, Florida and Japan.
998:binding which is essential for muscle cells.
2940:. Pennsylvania State University. 2012-05-02
2436:
2348:International Journal of Molecular Sciences
2205:
1974:
1911:
1909:
1907:
1888:. News » Science » Microbiology.
1757:
1755:
1753:
2670:
2217:"The zombie ants parasitized by the fungi
1943:11370/e6374602-b2a0-496c-b78e-774b34fb152b
1812:
1688:
1567:
1565:
1563:
986:and sarcoplasmic reticulum which provides
967:Some studies identified another compound,
710:
42:
3216:
3197:
3187:
3144:
3101:
3091:
3002:
2992:
2913:
2610:
2587:
2577:
2517:
2468:
2419:
2369:
2359:
2285:
2244:
2188:
2114:
2104:
2021:
2011:
1941:
1853:
1795:
1733:
1723:
1671:
1661:
1614:
1597:
302:After years of research, the taxonomy of
294:, anti-infective, and anticancer agents.
1904:
1879:
1877:
1875:
1873:
1750:
1289:Some studies proposed a theory in which
880:
738:core clade morphological characteristics
650:
2863:Journal of Traditional Chinese Medicine
2442:
2259:
2214:
2047:"Cordyceps: attack of the killer fungi"
1834:Communicative & Integrative Biology
1560:
1207:
233:, discovered by the British naturalist
14:
3510:
3050:
2886:Applied and Environmental Microbiology
2772:
2701:
2229:): new occurrence and natural history"
2156:
2131:
1883:
1488:
1242:
1080:
965:higher levels of sphingosine and GBA.
762:subclade morphological characteristics
400:Ophiocordyceps unilateralis sensu lato
3263:
3262:
1870:
1470:International Mycological Association
1284:
1251:
1115:species within Japanese cicadas, the
814:, using the evolutionary trait of an
787:In tropical forests, the ant species
663:
3025:
2604:
1452:
1215:suffers from an unidentified fungal
948:, combined with mechanical pressure.
683:species complex have both a sexual (
377:species forming a sister group with
2857:Wang Y, Enlai DA, Zhong JI (2013).
2408:Integrative and Comparative Biology
876:
795:spores. The spores attach to their
659:growing out of an infected host ant
339:was only recently brought forward.
24:
3066:
3051:Geddes, Linda (10 February 2023).
2645:
2045:Attenborough D (3 November 2008).
1316:
1281:, increasing the fungus' fitness.
1223:being viable, limiting the damage
1007:Natural products are host specific
25:
3574:
3237:
2443:Mangold, Colleen (17 July 2019).
2219:Ophiocordyceps camponotiatricipis
1534:Journal of Invertebrate Pathology
1200:, cholesterol medication such as
1172:conditions and light and are not
952:Convulsions and climbing behavior
930:spores onto the ant's exoskeleton
2746:Canadian Journal of Microbiology
2266:Proceedings. Biological Sciences
1776:Fungal Systematics and Evolution
842:quickly sets in, destroying the
306:is becoming increasingly clear.
63:
3044:
3019:
2930:
2850:
2736:
2646:Lu, Jennifer (April 18, 2019).
2639:
2613:Recent Patents on Biotechnology
2485:
2449:Journal of Experimental Biology
2302:
2253:
2038:
1403:and its prequel novel entitled
497:core clade as described in 2018
1482:
1187:
1057:
443:More recently in 2018, 15 new
297:
13:
1:
3250:.org. Accessed on 2010-08-22.
2824:Accounts of Chemical Research
2695:10.1016/S0031-9422(99)00272-1
1460:"Ophiocordyceps unilateralis"
1446:
1358:
1192:In 2009, a study showed that
782:
646:
629:subclade as described in 2018
432:of the fungi, leading to its
3125:G3: Genes, Genomes, Genetics
2994:10.1371/journal.pone.0036352
2579:10.1371/journal.pone.0004835
1725:10.1371/journal.pone.0017024
1663:10.1016/j.simyco.2017.12.002
1092:fungi produce various known
655:Schematic representation of
569:O. camponoti-novogranadensis
454:species associated with ants
407:Ophiocordyceps unilateralis
7:
3553:Suicide-inducing parasitism
3303:Ophiocordyceps unilateralis
3273:Ophiocordyceps unilateralis
3244:Ophiocordyceps unilateralis
3221:. Random House Publishing.
2260:Doherty JF (October 2020).
1884:Sample I (18 August 2010).
1766:Ophiocordyceps unilateralis
1489:Zimmer, Carl (2019-10-24).
1432:
1427:Ophiocordyceps unilateralis
1413:Ophiocordyceps unilateralis
1396:The Girl With All The Gifts
1381:Ophiocordyceps unilateralis
1371:Ophiocordyceps unilateralis
1048:Ophiocordyceps unilateralis
1026:Ophiocordyceps unilateralis
657:Ophiocordyceps unilateralis
389:Ophiocordyceps unilateralis
304:Ophiocordyceps unilateralis
275:Ophiocordyceps unilateralis
222:Ophiocordyceps unilateralis
169:Ophiocordyceps unilateralis
53:Ophiocordyceps unilateralis
36:Ophiocordyceps unilateralis
10:
3579:
2625:10.2174/187220807780809454
1264:The extended phenotype of
1127:Naphthoquinone derivatives
1032:since it occurs mainly in
982:levels due to the lack of
911:. Studies have shown that
544:O. camponoti-hippocrepidis
3451:
3271:
3093:10.1186/s12864-015-1812-x
2793:10.1007/s10295-005-0213-6
2722:10.1017/S0953756205003321
2310:"Rise of the zombie ants"
2106:10.1186/s12862-014-0166-3
1546:10.1016/j.jip.2012.08.007
1363:In the video game series
960:Two candidate compounds,
195:
188:
165:
158:
60:Scientific classification
58:
50:
41:
34:
2246:10.5943/mycosphere/8/9/1
2093:BMC Evolutionary Biology
687:) and an asexual morph (
679:Most species within the
529:O. camponoti-chartificis
387:which was later renamed
231:insect-pathogenic fungus
51:Dead ants infected with
3543:Mind-altering parasites
3518:Fungi described in 1865
3189:10.1073/pnas.1711673114
2013:10.1186/1472-6785-11-13
1922:The American Naturalist
1788:10.3114/fuse.2018.01.02
1405:The Boy on the Bridge -
1321:The principal hosts of
1121:Ophiocordyceps sinensis
711:Morphological variation
564:O. camponoti-nidulantis
559:O. camponoti- melanotic
481:subclade specialise on
414:complex of many species
3533:Fungi of South America
2679:Cordyceps unilateralis
2510:10.1098/rsbl.2010.0521
2278:10.1098/rspb.2020.1081
2181:10.1098/rsbl.2010.0521
1590:10.3114/sim.2007.57.01
1182:chemotherapeutic drugs
1050:was discovered in the
1024:Many studies describe
956:central nervous system
889:
852:sarcoplasmic reticular
660:
604:O. polyrhachis-furcata
579:O. camponoti-saundersi
539:O. camponoti-floridani
524:O. camponoti-bispinosi
514:O. camponoti-atricipis
430:reproductive isolation
385:Cordyceps unilateralis
205:Cordyceps unilateralis
3453:Torrubia unilateralis
3137:10.1534/g3.120.401290
1094:secondary metabolites
920:manipulates the ant.
884:
654:
574:O. camponoti-renggeri
554:O. camponoti-leonardi
534:O. camponoti-femorati
235:Alfred Russel Wallace
198:Torrubia unilateralis
3538:Ophiocordycipitaceae
3217:Sheldrake M (2020).
2906:10.1128/AEM.02744-08
2710:Mycological Research
2361:10.3390/ijms20184589
2227:Ophiocordycipitaceae
1770:Ophiocordycipitaceae
1401:2016 film adaptation
1295:O. unilateralis s.l.
1208:Fungal hyperparasite
681:O. unilateralis s.l.
549:O. camponoti-indiani
519:O. camponoti-balzani
279:secondary metabolism
225:, commonly known as
151:O. unilateralis
127:Ophiocordycipitaceae
3180:2017PNAS..11412590F
3174:(47): 12590–12595.
3032:Scientific American
3026:Parshall, Allison.
2985:2012PLoSO...736352A
2898:2009ApEnM..75.3721A
2652:National Geographic
2570:2009PLoSO...4.4835P
2314:www.natureindex.com
2215:Sobczak JF (2017).
1716:2011PLoSO...617024E
1650:Studies in Mycology
1578:Studies in Mycology
1440:Massospora cicadina
1344:Camponotus leonardi
1243:Parasite adaptation
1106:hypocholesterolemic
1081:Medicinal potential
805:Camponotus leonardi
789:Camponotus leonardi
625:Species within the
493:Species within the
456:which they termed "
452:mainly composed of
2461:10.1242/jeb.200683
2421:10.1093/icb/icu063
2272:(1936): 20201081.
1495:The New York Times
1393:In the 2014 novel
1285:Somatic investment
1252:Extended phenotype
1152:assays related to
890:
816:extended phenotype
664:Typical morphology
661:
3505:
3504:
3426:Open Tree of Life
3265:Taxon identifiers
3228:978-0-525-51031-4
2836:10.1021/ar040247r
2658:on April 18, 2019
2455:(14): jeb200683.
1846:10.4161/cib.16721
1275:natural selection
1154:antimalarial drug
1041:temperate forests
932:: The first step
829:temperate forests
725:ecological niches
460:core clade" and "
350:within the order
227:zombie-ant fungus
218:
217:
213:
202:
27:Species of fungus
18:Zombie ant fungus
16:(Redirected from
3570:
3498:
3497:
3485:
3484:
3472:
3471:
3470:
3444:
3443:
3434:
3433:
3421:
3420:
3408:
3407:
3395:
3394:
3382:
3381:
3369:
3368:
3356:
3355:
3343:
3342:
3330:
3329:
3317:
3316:
3307:
3306:
3305:
3292:
3291:
3290:
3260:
3259:
3232:
3211:
3201:
3191:
3158:
3148:
3131:(7): 2275–2296.
3115:
3105:
3095:
3061:
3060:
3048:
3042:
3041:
3039:
3038:
3023:
3017:
3016:
3006:
2996:
2964:
2949:
2948:
2946:
2945:
2934:
2928:
2927:
2917:
2877:
2871:
2870:
2854:
2848:
2847:
2819:
2813:
2812:
2776:
2770:
2769:
2740:
2734:
2733:
2716:(Pt 8): 936–40.
2705:
2699:
2698:
2674:
2668:
2667:
2665:
2663:
2654:. Archived from
2643:
2637:
2636:
2608:
2602:
2601:
2591:
2581:
2549:
2532:
2531:
2521:
2489:
2483:
2482:
2472:
2440:
2434:
2433:
2423:
2399:
2384:
2383:
2373:
2363:
2339:
2324:
2323:
2321:
2320:
2306:
2300:
2299:
2289:
2257:
2251:
2250:
2248:
2239:(9): 1261–1266.
2212:
2203:
2202:
2192:
2160:
2154:
2153:
2151:
2150:
2145:. 18 August 2010
2135:
2129:
2128:
2118:
2108:
2084:
2065:
2064:
2062:
2061:
2042:
2036:
2035:
2025:
2015:
1991:
1972:
1971:
1945:
1913:
1902:
1901:
1899:
1898:
1881:
1868:
1867:
1857:
1825:
1810:
1809:
1799:
1759:
1748:
1747:
1737:
1727:
1695:
1686:
1685:
1675:
1665:
1641:
1612:
1611:
1601:
1569:
1558:
1557:
1528:
1513:
1512:
1510:
1509:
1486:
1480:
1479:
1477:
1476:
1456:
1422:Cult of the Lamb
1333:
1075:
1028:distribution as
893:O. unilateralis'
877:Natural products
825:
768:O. kniphofioides
760:O. kniphofioides
641:O. kniphofioides
627:O. kniphofioides
594:O. naomipierceae
589:O. kimflemingiae
584:O. halabalaensis
509:O. blakebarnesii
479:O. kniphofioides
462:O. kniphofioides
292:immunomodulatory
284:natural products
253:, with the full
207:
200:
171:
68:
67:
46:
32:
31:
21:
3578:
3577:
3573:
3572:
3571:
3569:
3568:
3567:
3548:Parasitic fungi
3523:Fungi of Africa
3508:
3507:
3506:
3501:
3493:
3488:
3480:
3475:
3466:
3465:
3460:
3447:
3439:
3437:
3429:
3424:
3416:
3411:
3403:
3398:
3390:
3385:
3377:
3372:
3364:
3359:
3351:
3346:
3338:
3333:
3325:
3320:
3312:
3310:
3301:
3300:
3295:
3286:
3285:
3280:
3267:
3240:
3235:
3229:
3069:
3067:Further reading
3064:
3049:
3045:
3036:
3034:
3024:
3020:
2965:
2952:
2943:
2941:
2936:
2935:
2931:
2892:(11): 3721–32.
2878:
2874:
2855:
2851:
2820:
2816:
2777:
2773:
2758:10.1139/W11-043
2741:
2737:
2706:
2702:
2675:
2671:
2661:
2659:
2644:
2640:
2609:
2605:
2550:
2535:
2498:Biology Letters
2490:
2486:
2441:
2437:
2400:
2387:
2340:
2327:
2318:
2316:
2308:
2307:
2303:
2258:
2254:
2213:
2206:
2169:Biology Letters
2161:
2157:
2148:
2146:
2137:
2136:
2132:
2085:
2068:
2059:
2057:
2043:
2039:
1992:
1975:
1914:
1905:
1896:
1894:
1882:
1871:
1826:
1813:
1760:
1751:
1696:
1689:
1642:
1615:
1570:
1561:
1529:
1516:
1507:
1505:
1487:
1483:
1474:
1472:
1458:
1457:
1453:
1449:
1435:
1419:The video game
1407:all written by
1392:
1361:
1348:O. unilateralis
1337:O. unilateralis
1331:
1323:O. unilateralis
1319:
1317:Host adaptation
1310:O. unilateralis
1291:O. unilateralis
1287:
1271:O. unilateralis
1266:O. unilateralis
1259:O. unilateralis
1254:
1245:
1236:O. unilateralis
1229:O. unilateralis
1225:O. unilateralis
1213:O. unilateralis
1210:
1194:O. unilateralis
1190:
1161:O. unilateralis
1140:O. unilateralis
1136:O. unilateralis
1129:
1090:O. unilateralis
1083:
1073:
1064:O. unilateralis
1060:
1034:tropical forest
1022:
1013:O. unilateralis
1009:
1001:
934:O. unilateralis
928:O. unilateralis
918:O. unilateralis
913:O. unilateralis
909:immune response
887:O. unilateralis
879:
867:fruiting bodies
823:
793:O. unilateralis
785:
764:
744:O. unilateralis
740:
736:O. unilateralis
716:O. unilateralis
713:
670:O. unilateralis
666:
649:
504:O. albacongiuae
495:O. unilateralis
458:O. unilateralis
445:O. unilateralis
426:O. unilateralis
418:O. unilateralis
403:
348:Clavicipitaceae
318:
300:
269:O. unilateralis
265:fruiting bodies
243:O. unilateralis
239:tropical forest
203:
184:
173:
167:
154:
107:Sordariomycetes
62:
28:
23:
22:
15:
12:
11:
5:
3576:
3566:
3565:
3563:Fungus species
3560:
3555:
3550:
3545:
3540:
3535:
3530:
3525:
3520:
3503:
3502:
3500:
3499:
3486:
3473:
3457:
3455:
3449:
3448:
3446:
3445:
3435:
3422:
3409:
3396:
3383:
3370:
3357:
3344:
3331:
3318:
3308:
3293:
3277:
3275:
3269:
3268:
3257:
3256:
3251:
3239:
3238:External links
3236:
3234:
3233:
3227:
3214:
3212:
3159:
3116:
3070:
3068:
3065:
3063:
3062:
3043:
3018:
2950:
2929:
2872:
2849:
2830:(10): 813–23.
2814:
2771:
2735:
2700:
2689:(3): 453–457.
2683:Phytochemistry
2669:
2638:
2603:
2533:
2484:
2435:
2385:
2325:
2301:
2252:
2204:
2155:
2130:
2066:
2037:
1973:
1952:10.1086/603640
1934:10.1086/603640
1903:
1869:
1840:(5): 598–602.
1811:
1749:
1687:
1613:
1559:
1514:
1481:
1450:
1448:
1445:
1444:
1443:
1434:
1431:
1411:, a strain of
1385:climate change
1376:The Last of Us
1366:The Last of Us
1360:
1357:
1329:O.unilateralis
1318:
1315:
1286:
1283:
1253:
1250:
1244:
1241:
1209:
1206:
1189:
1186:
1132:Naphthoquinone
1128:
1125:
1117:Ophiocordyceps
1113:Ophiocordyceps
1086:Ophiocordyceps
1082:
1079:
1071:O.unilateralis
1059:
1056:
1021:
1018:
1008:
1005:
1000:
999:
972:
949:
926:Attachment of
922:
878:
875:
801:Ophiocordyceps
784:
781:
763:
757:
739:
733:
712:
709:
693:Ophiocordyceps
665:
662:
648:
645:
644:
643:
638:
622:
621:
616:
611:
606:
601:
596:
591:
586:
581:
576:
571:
566:
561:
556:
551:
546:
541:
536:
531:
526:
521:
516:
511:
506:
438:Ophiocordyceps
402:
397:
364:Ophiocordyceps
337:Ophiocordyceps
332:Ophiocordyceps
317:
315:Ophiocordyceps
308:
299:
296:
216:
215:
193:
192:
186:
185:
174:
163:
162:
156:
155:
148:
146:
142:
141:
138:Ophiocordyceps
134:
130:
129:
124:
120:
119:
114:
110:
109:
104:
100:
99:
94:
90:
89:
84:
80:
79:
74:
70:
69:
56:
55:
48:
47:
39:
38:
26:
9:
6:
4:
3:
2:
3575:
3564:
3561:
3559:
3556:
3554:
3551:
3549:
3546:
3544:
3541:
3539:
3536:
3534:
3531:
3529:
3528:Fungi of Asia
3526:
3524:
3521:
3519:
3516:
3515:
3513:
3496:
3491:
3487:
3483:
3478:
3474:
3469:
3463:
3459:
3458:
3456:
3454:
3450:
3442:
3436:
3432:
3427:
3423:
3419:
3414:
3410:
3406:
3401:
3397:
3393:
3388:
3384:
3380:
3375:
3371:
3367:
3362:
3358:
3354:
3349:
3345:
3341:
3336:
3332:
3328:
3323:
3319:
3315:
3309:
3304:
3298:
3294:
3289:
3283:
3279:
3278:
3276:
3274:
3270:
3266:
3261:
3255:
3252:
3249:
3245:
3242:
3241:
3230:
3224:
3220:
3215:
3213:
3209:
3205:
3200:
3195:
3190:
3185:
3181:
3177:
3173:
3169:
3165:
3160:
3156:
3152:
3147:
3142:
3138:
3134:
3130:
3126:
3122:
3117:
3113:
3109:
3104:
3099:
3094:
3089:
3085:
3081:
3077:
3072:
3071:
3058:
3054:
3047:
3033:
3029:
3022:
3014:
3010:
3005:
3000:
2995:
2990:
2986:
2982:
2979:(5): e36352.
2978:
2974:
2970:
2963:
2961:
2959:
2957:
2955:
2939:
2933:
2925:
2921:
2916:
2911:
2907:
2903:
2899:
2895:
2891:
2887:
2883:
2876:
2868:
2864:
2860:
2853:
2845:
2841:
2837:
2833:
2829:
2825:
2818:
2810:
2806:
2802:
2798:
2794:
2790:
2787:(4): 135–40.
2786:
2782:
2775:
2767:
2763:
2759:
2755:
2752:(8): 680–92.
2751:
2747:
2739:
2731:
2727:
2723:
2719:
2715:
2711:
2704:
2696:
2692:
2688:
2684:
2680:
2673:
2657:
2653:
2649:
2642:
2634:
2630:
2626:
2622:
2619:(2): 123–37.
2618:
2614:
2607:
2599:
2595:
2590:
2585:
2580:
2575:
2571:
2567:
2563:
2559:
2555:
2548:
2546:
2544:
2542:
2540:
2538:
2529:
2525:
2520:
2515:
2511:
2507:
2503:
2499:
2495:
2488:
2480:
2476:
2471:
2466:
2462:
2458:
2454:
2450:
2446:
2439:
2431:
2427:
2422:
2417:
2414:(2): 166–76.
2413:
2409:
2405:
2398:
2396:
2394:
2392:
2390:
2381:
2377:
2372:
2367:
2362:
2357:
2353:
2349:
2345:
2338:
2336:
2334:
2332:
2330:
2315:
2311:
2305:
2297:
2293:
2288:
2283:
2279:
2275:
2271:
2267:
2263:
2256:
2247:
2242:
2238:
2234:
2230:
2228:
2224:
2220:
2211:
2209:
2200:
2196:
2191:
2186:
2182:
2178:
2174:
2170:
2166:
2159:
2144:
2143:Science Daily
2140:
2134:
2126:
2122:
2117:
2112:
2107:
2102:
2098:
2094:
2090:
2083:
2081:
2079:
2077:
2075:
2073:
2071:
2056:
2055:BBC Worldwide
2052:
2048:
2041:
2033:
2029:
2024:
2019:
2014:
2009:
2005:
2001:
1997:
1990:
1988:
1986:
1984:
1982:
1980:
1978:
1969:
1965:
1961:
1957:
1953:
1949:
1944:
1939:
1935:
1931:
1928:(3): 424–33.
1927:
1923:
1919:
1912:
1910:
1908:
1893:
1892:
1887:
1880:
1878:
1876:
1874:
1865:
1861:
1856:
1851:
1847:
1843:
1839:
1835:
1831:
1824:
1822:
1820:
1818:
1816:
1807:
1803:
1798:
1793:
1789:
1785:
1781:
1777:
1773:
1771:
1767:
1758:
1756:
1754:
1745:
1741:
1736:
1731:
1726:
1721:
1717:
1713:
1710:(3): e17024.
1709:
1705:
1701:
1694:
1692:
1683:
1679:
1674:
1669:
1664:
1659:
1655:
1651:
1647:
1640:
1638:
1636:
1634:
1632:
1630:
1628:
1626:
1624:
1622:
1620:
1618:
1609:
1605:
1600:
1595:
1591:
1587:
1583:
1579:
1575:
1568:
1566:
1564:
1555:
1551:
1547:
1543:
1540:(3): 217–24.
1539:
1535:
1527:
1525:
1523:
1521:
1519:
1504:
1500:
1496:
1492:
1485:
1471:
1467:
1466:
1461:
1455:
1451:
1442:
1441:
1437:
1436:
1430:
1428:
1424:
1423:
1417:
1414:
1410:
1406:
1402:
1398:
1397:
1390:
1386:
1382:
1378:
1377:
1372:
1368:
1367:
1356:
1353:
1349:
1345:
1340:
1338:
1330:
1326:
1324:
1314:
1311:
1307:
1302:
1301:and dryness.
1300:
1296:
1292:
1282:
1280:
1276:
1272:
1267:
1262:
1260:
1249:
1240:
1237:
1232:
1230:
1226:
1222:
1218:
1217:hyperparasite
1214:
1205:
1203:
1199:
1195:
1185:
1183:
1179:
1175:
1171:
1167:
1162:
1157:
1155:
1151:
1150:
1145:
1141:
1137:
1133:
1124:
1122:
1118:
1114:
1109:
1107:
1103:
1099:
1095:
1091:
1087:
1078:
1072:
1067:
1065:
1055:
1053:
1049:
1044:
1042:
1037:
1035:
1031:
1027:
1017:
1014:
1004:
997:
993:
989:
985:
980:
976:
973:
970:
966:
963:
957:
953:
950:
947:
943:
939:
935:
931:
929:
924:
923:
921:
919:
914:
910:
904:
900:
898:
894:
888:
883:
874:
871:
868:
864:
860:
855:
853:
849:
845:
841:
837:
832:
830:
819:
817:
813:
808:
806:
802:
798:
794:
790:
780:
777:
773:
769:
761:
756:
754:
749:
745:
737:
732:
730:
726:
722:
717:
708:
706:
705:
700:
699:
694:
690:
686:
682:
677:
675:
671:
658:
653:
642:
639:
637:
634:
633:
632:
630:
628:
620:
617:
615:
612:
610:
607:
605:
602:
600:
597:
595:
592:
590:
587:
585:
582:
580:
577:
575:
572:
570:
567:
565:
562:
560:
557:
555:
552:
550:
547:
545:
542:
540:
537:
535:
532:
530:
527:
525:
522:
520:
517:
515:
512:
510:
507:
505:
502:
501:
500:
498:
496:
490:
488:
484:
480:
475:
471:
465:
463:
459:
455:
451:
446:
441:
439:
435:
431:
427:
421:
419:
415:
411:
410:
401:
396:
394:
390:
386:
382:
381:
380:Tolypocladium
376:
371:
369:
365:
361:
357:
353:
349:
345:
340:
338:
334:
333:
328:
327:
323:
316:
312:
307:
305:
295:
293:
289:
285:
280:
276:
272:
270:
266:
261:
256:
252:
249:of the tribe
248:
244:
240:
236:
232:
228:
224:
223:
214:
211:
206:
199:
194:
191:
187:
182:
178:
172:
170:
164:
161:
160:Binomial name
157:
153:
152:
147:
144:
143:
140:
139:
135:
132:
131:
128:
125:
122:
121:
118:
115:
112:
111:
108:
105:
102:
101:
98:
95:
92:
91:
88:
85:
82:
81:
78:
75:
72:
71:
66:
61:
57:
54:
49:
45:
40:
37:
33:
30:
19:
3452:
3272:
3218:
3171:
3167:
3128:
3124:
3083:
3080:BMC Genomics
3079:
3057:The Guardian
3056:
3046:
3035:. Retrieved
3031:
3021:
2976:
2972:
2942:. Retrieved
2932:
2889:
2885:
2875:
2866:
2862:
2852:
2827:
2823:
2817:
2784:
2780:
2774:
2749:
2745:
2738:
2713:
2709:
2703:
2686:
2682:
2678:
2672:
2662:December 24,
2660:. Retrieved
2656:the original
2651:
2641:
2616:
2612:
2606:
2564:(3): e4835.
2561:
2557:
2504:(1): 67–70.
2501:
2497:
2487:
2452:
2448:
2438:
2411:
2407:
2354:(18): 4589.
2351:
2347:
2317:. Retrieved
2313:
2304:
2269:
2265:
2255:
2236:
2232:
2226:
2222:
2218:
2175:(1): 67–70.
2172:
2168:
2158:
2147:. Retrieved
2133:
2096:
2092:
2058:. Retrieved
2051:Planet Earth
2050:
2040:
2003:
1999:
1925:
1921:
1895:. Retrieved
1891:The Guardian
1889:
1837:
1833:
1779:
1775:
1769:
1765:
1707:
1703:
1653:
1649:
1581:
1577:
1537:
1533:
1506:. Retrieved
1494:
1484:
1473:. Retrieved
1463:
1454:
1438:
1426:
1421:
1418:
1412:
1404:
1394:
1380:
1379:on HBO Max,
1374:
1370:
1364:
1362:
1351:
1350:is present,
1347:
1343:
1341:
1336:
1328:
1327:
1322:
1320:
1309:
1303:
1299:UV radiation
1294:
1290:
1288:
1277:acts on its
1270:
1265:
1263:
1258:
1255:
1246:
1235:
1233:
1228:
1224:
1212:
1211:
1193:
1191:
1160:
1158:
1147:
1143:
1139:
1135:
1130:
1120:
1116:
1112:
1110:
1102:hypoglycemic
1089:
1085:
1084:
1070:
1068:
1063:
1061:
1047:
1045:
1038:
1025:
1023:
1012:
1010:
1002:
974:
969:hypoxanthine
959:
951:
933:
927:
925:
917:
912:
905:
901:
896:
892:
891:
886:
872:
856:
848:mitochondria
833:
820:
809:
804:
800:
797:exoskeletons
792:
788:
786:
775:
767:
765:
759:
752:
747:
743:
741:
735:
715:
714:
704:Hymenostilbe
702:
696:
692:
680:
678:
669:
667:
656:
640:
635:
626:
624:
623:
618:
613:
609:O. pulvinata
608:
603:
598:
593:
588:
583:
578:
573:
568:
563:
558:
553:
548:
543:
538:
533:
528:
523:
518:
513:
508:
503:
494:
492:
491:
486:
478:
466:
461:
457:
444:
442:
437:
425:
422:
417:
406:
404:
399:
388:
384:
378:
374:
372:
368:monophyletic
363:
359:
343:
341:
336:
330:
324:
319:
314:
310:
303:
301:
274:
273:
268:
255:pathogenesis
242:
241:ecosystems.
226:
221:
220:
219:
204:
197:
196:
168:
166:
150:
149:
137:
52:
35:
29:
3400:NatureServe
3374:iNaturalist
3297:Wikispecies
2223:Hypocreales
2000:BMC Ecology
1656:: 119–160.
1584:(1): 5–59.
1389:Craig Mazin
1352:C. leonardi
1188:Polyketides
1058:Host impact
1054:(Germany).
1030:pantropical
1011:Effects of
962:sphingosine
897:C. leonardi
753:Camponotini
487:Camponotini
483:neotropical
464:subclade."
352:Hypocreales
298:Systematics
251:Camponotini
201:Tul. (1865)
117:Hypocreales
3512:Categories
3086:(1): 620.
3037:2023-02-11
2944:2013-03-04
2319:2021-01-19
2233:Mycosphere
2149:2010-09-12
2099:(1): 166.
2060:2013-04-21
1897:2010-08-22
1508:2022-04-30
1475:2011-07-19
1447:References
1409:M.R. Carey
1359:In fiction
1052:Messel pit
975:Death grip
836:Messel Pit
783:Life cycle
776:Hirsutella
748:Hirsutella
721:subspecies
698:Hirsutella
685:teleomorph
647:Morphology
599:O. ootakii
434:speciation
409:sensu lato
342:The genus
288:polyketide
97:Ascomycota
93:Division:
3468:Q64691835
3405:2.1072941
2006:(1): 13.
1782:: 13–22.
1503:0362-4331
1221:sporangia
1202:compactin
1174:cytotoxic
1108:targets.
1098:antitumor
938:chitinase
844:sarcomere
636:O. daceti
489:species.
474:septation
375:Cordyceps
360:Cordyceps
344:Cordyceps
326:Cordyceps
311:Cordyceps
260:mandibles
145:Species:
83:Kingdom:
77:Eukaryota
3490:MycoBank
3477:Fungorum
3462:Wikidata
3387:MycoBank
3348:Fungorum
3311:BioLib:
3282:Wikidata
3208:29114054
3155:32354705
3112:26285697
3013:22567151
2973:PLOS ONE
2924:19346345
2844:16231877
2809:22937549
2801:15891934
2766:21823977
2730:16175796
2633:19075836
2598:19279680
2558:PLOS ONE
2528:20719770
2479:31315924
2430:24907198
2380:31533250
2296:33049168
2199:20719770
2125:25085339
2032:21554670
1968:31283817
1960:19627240
1864:22046474
1806:32518897
1744:21399679
1704:PLOS ONE
1682:29910522
1608:18490993
1554:22959811
1465:MycoBank
1433:See also
1170:alkaline
1149:in vitro
946:protease
812:hemocoel
689:anamorph
674:pronotum
619:O. satoi
393:stromata
245:infects
229:, is an
190:Synonyms
123:Family:
73:Domain:
3441:1599624
3366:3450915
3288:Q127353
3248:UniProt
3199:5703306
3176:Bibcode
3146:7341126
3103:4545319
3004:3342268
2981:Bibcode
2915:2687288
2894:Bibcode
2589:2652714
2566:Bibcode
2519:3030878
2470:6679347
2371:6769763
2287:7657867
2190:3030878
2116:4174324
2023:3118224
1855:3204140
1797:7274273
1735:3047535
1712:Bibcode
1673:6002356
1599:2104736
1306:somatic
1198:patulin
988:calcium
979:leucine
863:mycelia
840:atrophy
729:fitness
614:O. rami
416:within
358:. When
208:(Tul.)
133:Genus:
113:Order:
103:Class:
3495:145303
3482:145303
3438:uBio:
3431:739625
3418:268505
3392:281145
3379:465287
3353:281145
3327:711976
3314:679032
3225:
3206:
3196:
3153:
3143:
3110:
3100:
3011:
3001:
2922:
2912:
2842:
2807:
2799:
2764:
2728:
2631:
2596:
2586:
2526:
2516:
2477:
2467:
2428:
2378:
2368:
2294:
2284:
2197:
2187:
2123:
2113:
2030:
2020:
1966:
1958:
1950:
1862:
1852:
1804:
1794:
1742:
1732:
1680:
1670:
1606:
1596:
1552:
1501:
1399:, its
1111:In an
1104:, and
996:myosin
942:lipase
859:hyphae
824:
772:ascoma
470:ascoma
450:clades
322:genera
212:(1883)
183:(1931)
3340:49WSK
2805:S2CID
1964:S2CID
1948:JSTOR
1332:'
1279:genes
1074:'
1062:When
992:actin
210:Sacc.
181:Petch
87:Fungi
3558:Ants
3413:NCBI
3361:GBIF
3322:BOLD
3223:ISBN
3204:PMID
3151:PMID
3108:PMID
3009:PMID
2920:PMID
2840:PMID
2797:PMID
2762:PMID
2726:PMID
2664:2022
2629:PMID
2594:PMID
2524:PMID
2475:PMID
2426:PMID
2376:PMID
2292:PMID
2195:PMID
2121:PMID
2028:PMID
1956:PMID
1860:PMID
1802:PMID
1740:PMID
1678:PMID
1604:PMID
1550:PMID
1499:ISSN
1166:acid
990:for
944:and
850:and
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