966:: 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
641:
871:
33:
888:. 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.
54:
1258:
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,
843:. 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.
1055:-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).
1344:
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.
1257:
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
1245:
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
970:
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
953:
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
891:
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
1343:
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
895:
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
1065:
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
1004:
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
1404:
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
465:
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
1227:
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
767:
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
707:
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
815:
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
810:
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
904:
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
1418:. 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.
960:, 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.
1152:
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
270:
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
246:
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
1323:
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
1301:
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.
1520:
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".
858:
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.
739:
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
1236:
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.
456:
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
436:
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
2732:
Amnuaykanjanasin A, Panchanawaporn S, Chutrakul C, Tanticharoen M (August 2011). "Genes differentially expressed under naphthoquinone-producing conditions in the entomopathogenic fungus
Ophiocordyceps unilateralis".
1328:-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.
251:
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
412:
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
780:
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
925:
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
943:: 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
665:
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.
992:
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.
759:
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
2768:
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".
807:
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.
1085:, as well as several structurally uncharacterised substances. These natural products are reportedly being investigated as potential leads in discovery efforts toward immunomodulatory,
355:. 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
1376:, 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.
1058:
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
1372:
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
260:
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.
1135:-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
3041:
1165:, 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
854:
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
1297:
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,
2636:
1145:
discovery. In addition to having antimalarial activities, all six of these secondary metabolites have been demonstrated to have anticancer and antibacterial activities.
657:
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,
2393:"From behavior to mechanisms: an integrative approach to the manipulation by a parasitic fungus (Ophiocordyceps unilateralis s.l.) of its host ants (Camponotus spp.)"
425:. Following this, a study conducted in Brazil delimited, using morphological comparisons of the ascospores, germination processes, and asexual morphs, four different
1193:, and antifungal treatments. It has also been reported that polyketides have other therapeutic effects such as antitumor, antioxidant and antiaging activities.
811:
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
1314:
evolved adaptive behaviors able to limit the contact rate between uninfected susceptible hosts and infected hosts, thereby reducing the risk of transmission.
2697:
Wongsa P, Tasanatai K, Watts P, Hywel-Jones N (August 2005). "Isolation and in vitro cultivation of the insect pathogenic fungus
Cordyceps unilateralis".
1906:
1689:"Hidden diversity behind the zombie-ant fungus Ophiocordyceps unilateralis: four new species described from carpenter ants in Minas Gerais, Brazil"
796:; despite this, the fungus may parasitize other closely related species of ants with lesser degrees of host manipulation and reproductive success.
3016:
1286:
species as they can produce and release within the air, clear and thin-walled spores which are susceptible to environmental conditions such as
862:
The term "zombie ants" has been used in popular media as well as scientific articles, but has also been described as "catchy, yet misleading."
2666:
Kittakoopa P, Punyaa J, Kongsaeree P, Lertwerawat Y, Jintasirikul A, Tanticharoena M, Thebtaranonth Y (1999). "Bioactive naphthoquinones from
3401:
2871:"Insect-specific polyketide synthases (PKSs), potential PKS-nonribosomal peptide synthetase hybrids, and novel PKS clades in tropical fungi"
1066:
season stimulates fungal development, which leads to more spores being released and ultimately more individuals being infected and killed.
1035:
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
417:
species can only successfully infect and manipulate one host ant species. There is a possibility that this resulted in or reinforced the
2811:
Isaka M, Kittakoop P, Kirtikara K, Hywel-Jones NL, Thebtaranonth Y (October 2005). "Bioactive substances from insect pathogenic fungi".
1108:
replaces the symbiotic bacteria within the cicadas to help the host process sap as nutrients, unlike other related species, such as the
2298:
1819:"Ophiocordyceps unilateralis: A keystone species for unraveling ecosystem functioning and biodiversity of fungi in tropical forests?"
3153:"Three-dimensional visualization and a deep-learning model reveal complex fungal parasite networks in behaviorally manipulated ants"
2848:"A Retrospective Analysis of Cordyceps Anti-Tuberculosis Capsule Combined with Chemotherapy for 614 Cases of Secondary Tuberculosis"
3349:
1362:
has evolved to infect humans, thus creating zombie-like enemies in the game. Also, in episode two of the 2023 television series
640:
744:
species. Once the host is killed by the fungus, it is commonly found fixed through their mandibles onto the surfaces of leaves.
3065:"Gene expression during zombie ant biting behavior reflects the complexity underlying fungal parasitic behavioral manipulation"
1380:, who co-wrote and produced the series, said that everything the series suggests fungi do, they have done forever in real life.
2644:
3291:
3215:
1458:
461:(perithecia) growing from the stroma) and microscopic traits (e.g. the morphology of the ascospores in terms of size, shape,
3541:
816:
vegetative growth and sporulation either fails to occur or results in undersized and abnormal reproductive structures. In
1414:
features an ant character named Sozo who is implied to be under the influence of a parasitic fungus similar in nature to
788:
and eventually break through using mechanical pressure and enzymes. Like other fungi pathogenic to insects in the genus
324:. There have been many debates about whether the zombie-ant fungus (and other fungi) belonged to one or to the other as
1389:
2333:"Specialist and Generalist Fungal Parasites Induce Distinct Biochemical Changes in the Mandible Muscles of Their Host"
343:. The classification was based on different morphological characteristics such as filiform ascospores and cylindrical
680:). These are different in terms of their function and characteristics. Generally, the asexual morphs identified for
3531:
3506:
3406:
429:
species. Afterwards, three new species were described in the
Brazilian Amazon, six in Thailand, and one in Japan.
3521:
735:
core clade, as described in 2018, has distinct morphological characteristics. It exhibits a single stroma with a
2600:
Xiao JH, Zhong JJ (2007). "Secondary metabolites from
Cordyceps species and their antitumor activity studies".
1405:
influencing animals with a fungus that invades human brains and influences their actions toward its advantage.
1335:, provided evidence for the avoidance of the forest floor by the host ants as a defence method. In areas where
1874:
3526:
1282:
has another possible form of adaptation which ensures its repeated reproduction. This would be crucial for
1223:
The graveyards of dead ants are numerous and spread throughout the surrounding area of the colony. Though
799:
Yeast stages of the fungus spread in the ant's body and presumably produce compounds that affect the ant's
53:
712:
of ant can occur within the same area, which means that in order to coexist they have to occupy different
3315:
3310:
3110:"Genetic Underpinnings of Host Manipulation by Ophiocordyceps as Revealed by Comparative Transcriptomics"
1384:
248:
1123:
derivatives are an example of secondary metabolite with important pharmaceutical potentials produced by
3063:
de Bekker C, Ohm RA, Loreto RG, Sebastian A, Albert I, Merrow M, Brachmann A, Hughes DP (August 2015).
1364:
372:, into Ophiocordycipitaceae. Fungi able to parasitize ants were also included in the transfer, such as
1931:
1905:
Andersen SB, Gerritsma S, Yusah KM, Mayntz D, Hywel-Jones NL, Billen J, et al. (September 2009).
884:
life cycle includes and depends on the infection and the manipulation of a carpenter ant, principally
820:, the typical behavior of zombie ants is to attach themselves to the lower side of twigs, not leaves.
3470:
3341:
1479:
1220:
as the fungal stalk emerges from the ant's body, which can stop the stalk from releasing its spores.
947:(CNS), which enables it to manipulate the ant to reach the forest floor and climb up the vegetation.
380:. Following this study, multiple traits such as the production of darkly pigmented, hard to flexible
661:
typically generates a single, wiry yet pliant, darkly pigmented stroma which arises from the dorsal
3536:
3511:
3393:
1025:
ecosystems. However, there are some reports of the zombie-ant fungus in warm-temperate ecosystems.
1005:
heterogeneous nature of the fungus which secretes different metabolites according to host species.
874:
Schematic representation of the ant behavioral manipulation caused by natural products secreted by
870:
3551:
2127:
2076:
de Bekker C, Quevillon LE, Smith PB, Fleming KR, Ghosh D, Patterson AD, Hughes DP (August 2014).
219:
2926:
1262:
controls the ant's behavior and this manipulation represents an adaptation for the fungus where
3516:
3253:
3151:
Fredericksen MA, Zhang Y, Hazen ML, Loreto RG, Mangold CA, Chen DZ, Hughes DP (November 2017).
2869:
Amnuaykanjanasin A, Phonghanpot S, Sengpanich N, Cheevadhanarak S, Tanticharoen M (June 2009).
2847:
2543:"Graveyards on the move: the spatio-temporal distribution of dead ophiocordyceps-infected ants"
1170:
972:
944:
840:
418:
3042:"'A growing threat to human health': we are ill-equipped for the dangers of fungal infections"
1448:
1028:
Its distribution includes tropical rainforests located in Brazil, Australia and
Thailand, and
3429:
1985:"Behavioral mechanisms and morphological symptoms of zombie ants dying from fungal infection"
1185:
also produces polyketides. These secondary metabolites have been used in antibiotics such as
1112:, which is a traditional immune booster and cancer treatment in Tibetan and Chinese culture.
1082:
827:, which is 48 million years old. Once the mandibles of the ant are secured to the leaf vein,
223:
148:
3419:
1208:, reported in the lay press as the "antizombie-fungus fungus", that results in only 6–7% of
768:
usually die at the base of large trees in the
Amazonian rainforest, among the moss carpets.
716:. Consequently, the fungi may have evolved at the subspecies level in order to maximize its
362:
The new molecular phylogenetics studies contradicted the older classification and moved all
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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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2011:
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2016:
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1263:
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717:
713:
253:
2797:
1983:
Hughes DP, Andersen SB, Hywel-Jones NL, Himaman W, Billen J, Boomsma JJ (May 2011).
1956:
823:
A search of plant-fossil databases revealed similar marks on a fossil leaf from the
3328:
3208:
Entangled Life: How Fungi Make Our Worlds, Change Our Minds & Shape Our
Futures
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2006:
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Sung GH, Hywel-Jones NL, Sung JM, Luangsa-Ard JJ, Shrestha B, Spatafora JW (2007).
1530:
1409:
1029:
817:
280:
2541:
Pontoppidan MB, Himaman W, Hywel-Jones NL, Boomsma JJ, Hughes DP (12 March 2009).
3456:
2982:
2956:
Andersen SB, Ferrari M, Evans HC, Elliot SL, Boomsma JJ, Hughes DP (2 May 2012).
2567:
1713:
1651:
1022:
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families: the Clavicipitaceae, the Cordycipitaceae and the Ophiocordycipitaceae.
336:
272:
227:
95:
3465:
3336:
3157:
Proceedings of the National Academy of Sciences of the United States of America
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125:
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2781:
2710:
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2035:
1940:
1534:
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invade more soft tissues and structurally fortify the ant's exoskeleton. More
401:, which means 'in the broad sense', because the species actually represents a
309:
Throughout history there has been confusion about the distinction between the
32:
3546:
3500:
3276:
2131:
2043:
1491:
1205:
368:
275:
chemists, with corresponding discovery of small molecule agents (e.g. of the
3177:
2331:
Zheng S, Loreto R, Smith P, Patterson A, Hughes D, Wang L (September 2019).
2001:
1776:
896:
comes across an array of environments such as different host tissues or the
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3100:
3001:
2912:
2832:
2789:
2754:
2718:
2621:
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2113:
2020:
1948:
1879:
1852:
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1670:
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1542:
1294:
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1166:
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957:
836:
723:
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356:
243:
3302:
3232:
3125:
2078:"Species-specific ant brain manipulation by a specialized fungal parasite"
1907:"The life of a dead ant: the expression of an adaptive extended phenotype"
1228:
colony. Additional fungi also grant beneficial assistance to the colony.
1169:
patients, by improving symptoms and enhancing immunity when combined with
747:
3388:
3362:
3285:
2894:
2409:
2392:
2349:
1563:"Phylogenetic classification of Cordyceps and the clavicipitaceous fungi"
1377:
1077:
are known in the pharmaceutical world to be a medically-important group.
1018:
950:
785:
471:
340:
239:
105:
2481:
Hughes, David P.; Wappler, Torsten; Labandeira, Conrad C. (2011-02-23).
2251:"When fiction becomes fact: exaggerating host manipulation by parasites"
3354:
2449:
1209:
1040:
824:
709:
686:
673:
422:
276:
85:
3367:
3108:
Will I, Das B, Trinh T, Brachmann A, Ohm RA, de Bekker C (July 2020).
3017:"Could the Zombie Fungus in TV's The Last of Us Really Infect People?"
2824:
1834:
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1086:
1008:
926:
832:
462:
314:
169:
65:
3247:
2746:
2540:
384:
were defined as characteristics of the family Ophiocordycipitaceae.
351:
were first classified, there was no concrete evidence for the genus
256:
grow from the ant's head, rupturing to release the fungus's spores.
3478:
3450:
3375:
3270:
1922:
1453:
1158:
1137:
934:
800:
677:
662:
2810:
1250:
represents one of the best-known examples of extended phenotypes.
335:
comprises over 400 species, historically classified in the family
3236:
2434:"Zombie ant death grip due to hypercontracted mandibular muscles"
1480:"After This Fungus Turns Ants Into Zombies, Their Bodies Explode"
1186:
976:
967:
851:
828:
1982:
2696:
1216:
inflicts on ant colonies. The hyperparasite moves in to attack
1127:. Six known naphthoquinone derivatives have been isolated from
984:
930:
847:
760:
458:
438:
75:
2075:
1753:"Epitypification and re-description of the zombie-ant fungus,
1560:
2958:"Disease dynamics in a specialized parasite of ant societies"
1267:
1148:
Moreover, the use of red naphthoquinone pigments produced by
980:
344:
310:
3150:
2767:
2483:"Ancient death-grip leaf scars reveal ant–fungal parasitism"
2154:"Ancient death-grip leaf scars reveal ant-fungal parasitism"
1904:
1875:"'Zombie ants' controlled by parasitic fungus for 48m years"
3062:
1154:
846:
The fungus then kills the ant and continues to grow as its
2955:
2927:"The Zombie-Ant Fungus Is Under Attack, Research Reveals"
2330:
892:
protect the carcass from microbial and scavenger attack.
387:
235:
1751:
Evans HC, AraĂşjo JP, Halfeld VR, Hughes DP (June 2018).
1750:
1635:"Ophiocordyceps. I. Myrmecophilous hirsutelloid species"
3242:
2480:
2151:
2128:"Fossil Reveals 48-Million-Year History of Zombie Ants"
1632:
3243:
An Electronic Monograph of Cordyceps and Related Fungi
2770:
Journal of Industrial Microbiology & Biotechnology
2152:
Hughes DP, Wappler T, Labandeira CC (February 2011) .
1633:
AraĂşjo JP, Evans HC, Kepler R, Hughes DP (June 2018).
474:
ants, whereas species in the core clade specialise on
3107:
2390:
995:
394:
The fungus's scientific name is sometimes written as
266:
and related species are known to engage in an active
2862:
1131:, namely erythrostominone, deoxyerythrostominone, 4-
1515:
1513:
1511:
1509:
1507:
2637:"How a parasitic fungus turns ants into 'zombies'"
2536:
2534:
2532:
2530:
2528:
2526:
1009:Geographic distribution and first known appearance
835:connections in the muscle fibers and reducing the
1817:Evans HC, Elliot SL, Hughes DP (September 2011).
1816:
1686:
763:. Moreover, species within this subclade share a
3498:
2951:
2949:
2947:
2945:
2943:
2071:
2069:
2067:
2065:
2063:
2061:
2059:
1504:
2523:
2386:
2384:
2382:
2380:
2378:
2326:
2324:
2322:
2320:
2318:
279:family) of potential interest for use as human
226:in 1859, and currently found predominantly in
2845:
2391:de Bekker C, Merrow M, Hughes DP (July 2014).
2199:
2197:
1978:
1976:
1974:
1972:
1970:
1968:
1966:
1331:Moreover, one of the fungus' principal hosts,
792:, the fungus targets a specific host species,
708:species (host-specific infections). Different
298:
2940:
2056:
1812:
1810:
1808:
1806:
1804:
1687:Evans HC, Elliot SL, Hughes DP (March 2011).
1682:
1680:
1628:
1626:
1115:
696:, two genera of asexually reproducing fungi.
2804:
2375:
2315:
2033:
1624:
1622:
1620:
1618:
1616:
1614:
1612:
1610:
1608:
1606:
1032:found in South Carolina, Florida and Japan.
987:binding which is essential for muscle cells.
2929:. Pennsylvania State University. 2012-05-02
2425:
2337:International Journal of Molecular Sciences
2194:
1963:
1900:
1898:
1896:
1877:. News » Science » Microbiology.
1746:
1744:
1742:
2659:
2206:"The zombie ants parasitized by the fungi
1932:11370/e6374602-b2a0-496c-b78e-774b34fb152b
1801:
1677:
1556:
1554:
1552:
975:and sarcoplasmic reticulum which provides
956:Some studies identified another compound,
699:
31:
3205:
3186:
3176:
3133:
3090:
3080:
2991:
2981:
2902:
2599:
2576:
2566:
2506:
2457:
2408:
2358:
2348:
2274:
2233:
2177:
2103:
2093:
2010:
2000:
1930:
1842:
1784:
1722:
1712:
1660:
1650:
1603:
1586:
291:After years of research, the taxonomy of
283:, anti-infective, and anticancer agents.
1893:
1868:
1866:
1864:
1862:
1739:
1278:Some studies proposed a theory in which
869:
727:core clade morphological characteristics
639:
2852:Journal of Traditional Chinese Medicine
2431:
2248:
2203:
2036:"Cordyceps: attack of the killer fungi"
1823:Communicative & Integrative Biology
1549:
1196:
222:, discovered by the British naturalist
3499:
3039:
2875:Applied and Environmental Microbiology
2761:
2690:
2218:): new occurrence and natural history"
2145:
2120:
1872:
1477:
1231:
1069:
954:higher levels of sphingosine and GBA.
751:subclade morphological characteristics
389:Ophiocordyceps unilateralis sensu lato
3252:
3251:
1859:
1459:International Mycological Association
1273:
1240:
1104:species within Japanese cicadas, the
803:, using the evolutionary trait of an
776:In tropical forests, the ant species
652:
3014:
2593:
1441:
1204:suffers from an unidentified fungal
937:, combined with mechanical pressure.
672:species complex have both a sexual (
366:species forming a sister group with
2846:Wang Y, Enlai DA, Zhong JI (2013).
2397:Integrative and Comparative Biology
865:
784:spores. The spores attach to their
648:growing out of an infected host ant
328:was only recently brought forward.
13:
3055:
3040:Geddes, Linda (10 February 2023).
2634:
2034:Attenborough D (3 November 2008).
1305:
1270:, increasing the fungus' fitness.
1212:being viable, limiting the damage
996:Natural products are host specific
14:
3563:
3226:
2432:Mangold, Colleen (17 July 2019).
2208:Ophiocordyceps camponotiatricipis
1523:Journal of Invertebrate Pathology
1189:, cholesterol medication such as
1161:conditions and light and are not
941:Convulsions and climbing behavior
919:spores onto the ant's exoskeleton
2735:Canadian Journal of Microbiology
2255:Proceedings. Biological Sciences
1765:Fungal Systematics and Evolution
831:quickly sets in, destroying the
295:is becoming increasingly clear.
52:
3033:
3008:
2919:
2839:
2725:
2635:Lu, Jennifer (April 18, 2019).
2628:
2602:Recent Patents on Biotechnology
2474:
2438:Journal of Experimental Biology
2291:
2242:
2027:
1392:and its prequel novel entitled
486:core clade as described in 2018
1471:
1176:
1046:
432:More recently in 2018, 15 new
286:
1:
3239:.org. Accessed on 2010-08-22.
2813:Accounts of Chemical Research
2684:10.1016/S0031-9422(99)00272-1
1449:"Ophiocordyceps unilateralis"
1435:
1347:
1181:In 2009, a study showed that
771:
635:
618:subclade as described in 2018
421:of the fungi, leading to its
3114:G3: Genes, Genomes, Genetics
2983:10.1371/journal.pone.0036352
2568:10.1371/journal.pone.0004835
1714:10.1371/journal.pone.0017024
1652:10.1016/j.simyco.2017.12.002
1081:fungi produce various known
644:Schematic representation of
558:O. camponoti-novogranadensis
443:species associated with ants
396:Ophiocordyceps unilateralis
7:
3542:Suicide-inducing parasitism
3292:Ophiocordyceps unilateralis
3262:Ophiocordyceps unilateralis
3233:Ophiocordyceps unilateralis
3210:. Random House Publishing.
2249:Doherty JF (October 2020).
1873:Sample I (18 August 2010).
1755:Ophiocordyceps unilateralis
1478:Zimmer, Carl (2019-10-24).
1421:
1416:Ophiocordyceps unilateralis
1402:Ophiocordyceps unilateralis
1385:The Girl With All The Gifts
1370:Ophiocordyceps unilateralis
1360:Ophiocordyceps unilateralis
1037:Ophiocordyceps unilateralis
1015:Ophiocordyceps unilateralis
646:Ophiocordyceps unilateralis
378:Ophiocordyceps unilateralis
293:Ophiocordyceps unilateralis
264:Ophiocordyceps unilateralis
211:Ophiocordyceps unilateralis
158:Ophiocordyceps unilateralis
42:Ophiocordyceps unilateralis
25:Ophiocordyceps unilateralis
10:
3568:
2614:10.2174/187220807780809454
1253:The extended phenotype of
1116:Naphthoquinone derivatives
1021:since it occurs mainly in
971:levels due to the lack of
900:. Studies have shown that
533:O. camponoti-hippocrepidis
3440:
3260:
3082:10.1186/s12864-015-1812-x
2782:10.1007/s10295-005-0213-6
2711:10.1017/S0953756205003321
2299:"Rise of the zombie ants"
2095:10.1186/s12862-014-0166-3
1535:10.1016/j.jip.2012.08.007
1352:In the video game series
949:Two candidate compounds,
184:
177:
154:
147:
49:Scientific classification
47:
39:
30:
23:
2235:10.5943/mycosphere/8/9/1
2082:BMC Evolutionary Biology
676:) and an asexual morph (
668:Most species within the
518:O. camponoti-chartificis
376:which was later renamed
220:insect-pathogenic fungus
40:Dead ants infected with
3532:Mind-altering parasites
3507:Fungi described in 1865
3178:10.1073/pnas.1711673114
2002:10.1186/1472-6785-11-13
1911:The American Naturalist
1777:10.3114/fuse.2018.01.02
1394:The Boy on the Bridge -
1310:The principal hosts of
1110:Ophiocordyceps sinensis
700:Morphological variation
553:O. camponoti-nidulantis
548:O. camponoti- melanotic
470:subclade specialise on
403:complex of many species
3522:Fungi of South America
2668:Cordyceps unilateralis
2499:10.1098/rsbl.2010.0521
2267:10.1098/rspb.2020.1081
2170:10.1098/rsbl.2010.0521
1579:10.3114/sim.2007.57.01
1171:chemotherapeutic drugs
1039:was discovered in the
1013:Many studies describe
945:central nervous system
878:
841:sarcoplasmic reticular
649:
593:O. polyrhachis-furcata
568:O. camponoti-saundersi
528:O. camponoti-floridani
513:O. camponoti-bispinosi
503:O. camponoti-atricipis
419:reproductive isolation
374:Cordyceps unilateralis
194:Cordyceps unilateralis
3442:Torrubia unilateralis
3126:10.1534/g3.120.401290
1083:secondary metabolites
909:manipulates the ant.
873:
643:
563:O. camponoti-renggeri
543:O. camponoti-leonardi
523:O. camponoti-femorati
224:Alfred Russel Wallace
187:Torrubia unilateralis
3527:Ophiocordycipitaceae
3206:Sheldrake M (2020).
2895:10.1128/AEM.02744-08
2699:Mycological Research
2350:10.3390/ijms20184589
2216:Ophiocordycipitaceae
1759:Ophiocordycipitaceae
1390:2016 film adaptation
1284:O. unilateralis s.l.
1197:Fungal hyperparasite
670:O. unilateralis s.l.
538:O. camponoti-indiani
508:O. camponoti-balzani
268:secondary metabolism
214:, commonly known as
140:O. unilateralis
116:Ophiocordycipitaceae
3169:2017PNAS..11412590F
3163:(47): 12590–12595.
3021:Scientific American
3015:Parshall, Allison.
2974:2012PLoSO...736352A
2887:2009ApEnM..75.3721A
2641:National Geographic
2559:2009PLoSO...4.4835P
2303:www.natureindex.com
2204:Sobczak JF (2017).
1705:2011PLoSO...617024E
1639:Studies in Mycology
1567:Studies in Mycology
1429:Massospora cicadina
1333:Camponotus leonardi
1232:Parasite adaptation
1095:hypocholesterolemic
1070:Medicinal potential
794:Camponotus leonardi
778:Camponotus leonardi
614:Species within the
482:Species within the
445:which they termed "
441:mainly composed of
2450:10.1242/jeb.200683
2410:10.1093/icb/icu063
2261:(1936): 20201081.
1484:The New York Times
1382:In the 2014 novel
1274:Somatic investment
1241:Extended phenotype
1141:assays related to
879:
805:extended phenotype
653:Typical morphology
650:
3494:
3493:
3415:Open Tree of Life
3254:Taxon identifiers
3217:978-0-525-51031-4
2825:10.1021/ar040247r
2647:on April 18, 2019
2444:(14): jeb200683.
1835:10.4161/cib.16721
1264:natural selection
1143:antimalarial drug
1030:temperate forests
921:: The first step
818:temperate forests
714:ecological niches
449:core clade" and "
339:within the order
216:zombie-ant fungus
207:
206:
202:
191:
16:Species of fungus
3559:
3487:
3486:
3474:
3473:
3461:
3460:
3459:
3433:
3432:
3423:
3422:
3410:
3409:
3397:
3396:
3384:
3383:
3371:
3370:
3358:
3357:
3345:
3344:
3332:
3331:
3319:
3318:
3306:
3305:
3296:
3295:
3294:
3281:
3280:
3279:
3249:
3248:
3221:
3200:
3190:
3180:
3147:
3137:
3120:(7): 2275–2296.
3104:
3094:
3084:
3050:
3049:
3037:
3031:
3030:
3028:
3027:
3012:
3006:
3005:
2995:
2985:
2953:
2938:
2937:
2935:
2934:
2923:
2917:
2916:
2906:
2866:
2860:
2859:
2843:
2837:
2836:
2808:
2802:
2801:
2765:
2759:
2758:
2729:
2723:
2722:
2705:(Pt 8): 936–40.
2694:
2688:
2687:
2663:
2657:
2656:
2654:
2652:
2643:. Archived from
2632:
2626:
2625:
2597:
2591:
2590:
2580:
2570:
2538:
2521:
2520:
2510:
2478:
2472:
2471:
2461:
2429:
2423:
2422:
2412:
2388:
2373:
2372:
2362:
2352:
2328:
2313:
2312:
2310:
2309:
2295:
2289:
2288:
2278:
2246:
2240:
2239:
2237:
2228:(9): 1261–1266.
2201:
2192:
2191:
2181:
2149:
2143:
2142:
2140:
2139:
2134:. 18 August 2010
2124:
2118:
2117:
2107:
2097:
2073:
2054:
2053:
2051:
2050:
2031:
2025:
2024:
2014:
2004:
1980:
1961:
1960:
1934:
1902:
1891:
1890:
1888:
1887:
1870:
1857:
1856:
1846:
1814:
1799:
1798:
1788:
1748:
1737:
1736:
1726:
1716:
1684:
1675:
1674:
1664:
1654:
1630:
1601:
1600:
1590:
1558:
1547:
1546:
1517:
1502:
1501:
1499:
1498:
1475:
1469:
1468:
1466:
1465:
1445:
1411:Cult of the Lamb
1322:
1064:
1017:distribution as
882:O. unilateralis'
866:Natural products
814:
757:O. kniphofioides
749:O. kniphofioides
630:O. kniphofioides
616:O. kniphofioides
583:O. naomipierceae
578:O. kimflemingiae
573:O. halabalaensis
498:O. blakebarnesii
468:O. kniphofioides
451:O. kniphofioides
281:immunomodulatory
273:natural products
242:, with the full
196:
189:
160:
57:
56:
35:
21:
20:
3567:
3566:
3562:
3561:
3560:
3558:
3557:
3556:
3537:Parasitic fungi
3512:Fungi of Africa
3497:
3496:
3495:
3490:
3482:
3477:
3469:
3464:
3455:
3454:
3449:
3436:
3428:
3426:
3418:
3413:
3405:
3400:
3392:
3387:
3379:
3374:
3366:
3361:
3353:
3348:
3340:
3335:
3327:
3322:
3314:
3309:
3301:
3299:
3290:
3289:
3284:
3275:
3274:
3269:
3256:
3229:
3224:
3218:
3058:
3056:Further reading
3053:
3038:
3034:
3025:
3023:
3013:
3009:
2954:
2941:
2932:
2930:
2925:
2924:
2920:
2881:(11): 3721–32.
2867:
2863:
2844:
2840:
2809:
2805:
2766:
2762:
2747:10.1139/W11-043
2730:
2726:
2695:
2691:
2664:
2660:
2650:
2648:
2633:
2629:
2598:
2594:
2539:
2524:
2487:Biology Letters
2479:
2475:
2430:
2426:
2389:
2376:
2329:
2316:
2307:
2305:
2297:
2296:
2292:
2247:
2243:
2202:
2195:
2158:Biology Letters
2150:
2146:
2137:
2135:
2126:
2125:
2121:
2074:
2057:
2048:
2046:
2032:
2028:
1981:
1964:
1903:
1894:
1885:
1883:
1871:
1860:
1815:
1802:
1749:
1740:
1685:
1678:
1631:
1604:
1559:
1550:
1518:
1505:
1496:
1494:
1476:
1472:
1463:
1461:
1447:
1446:
1442:
1438:
1424:
1408:The video game
1396:all written by
1381:
1350:
1337:O. unilateralis
1326:O. unilateralis
1320:
1312:O. unilateralis
1308:
1306:Host adaptation
1299:O. unilateralis
1280:O. unilateralis
1276:
1260:O. unilateralis
1255:O. unilateralis
1248:O. unilateralis
1243:
1234:
1225:O. unilateralis
1218:O. unilateralis
1214:O. unilateralis
1202:O. unilateralis
1199:
1183:O. unilateralis
1179:
1150:O. unilateralis
1129:O. unilateralis
1125:O. unilateralis
1118:
1079:O. unilateralis
1072:
1062:
1053:O. unilateralis
1049:
1023:tropical forest
1011:
1002:O. unilateralis
998:
990:
923:O. unilateralis
917:O. unilateralis
907:O. unilateralis
902:O. unilateralis
898:immune response
876:O. unilateralis
868:
856:fruiting bodies
812:
782:O. unilateralis
774:
753:
733:O. unilateralis
729:
725:O. unilateralis
705:O. unilateralis
702:
659:O. unilateralis
655:
638:
493:O. albacongiuae
484:O. unilateralis
447:O. unilateralis
434:O. unilateralis
415:O. unilateralis
407:O. unilateralis
392:
337:Clavicipitaceae
307:
289:
258:O. unilateralis
254:fruiting bodies
232:O. unilateralis
228:tropical forest
192:
173:
162:
156:
143:
96:Sordariomycetes
51:
17:
12:
11:
5:
3565:
3555:
3554:
3552:Fungus species
3549:
3544:
3539:
3534:
3529:
3524:
3519:
3514:
3509:
3492:
3491:
3489:
3488:
3475:
3462:
3446:
3444:
3438:
3437:
3435:
3434:
3424:
3411:
3398:
3385:
3372:
3359:
3346:
3333:
3320:
3307:
3297:
3282:
3266:
3264:
3258:
3257:
3246:
3245:
3240:
3228:
3227:External links
3225:
3223:
3222:
3216:
3203:
3201:
3148:
3105:
3059:
3057:
3054:
3052:
3051:
3032:
3007:
2939:
2918:
2861:
2838:
2819:(10): 813–23.
2803:
2760:
2724:
2689:
2678:(3): 453–457.
2672:Phytochemistry
2658:
2627:
2592:
2522:
2473:
2424:
2374:
2314:
2290:
2241:
2193:
2144:
2119:
2055:
2026:
1962:
1941:10.1086/603640
1923:10.1086/603640
1892:
1858:
1829:(5): 598–602.
1800:
1738:
1676:
1602:
1548:
1503:
1470:
1439:
1437:
1434:
1433:
1432:
1423:
1420:
1400:, a strain of
1374:climate change
1365:The Last of Us
1355:The Last of Us
1349:
1346:
1318:O.unilateralis
1307:
1304:
1275:
1272:
1242:
1239:
1233:
1230:
1198:
1195:
1178:
1175:
1121:Naphthoquinone
1117:
1114:
1106:Ophiocordyceps
1102:Ophiocordyceps
1075:Ophiocordyceps
1071:
1068:
1060:O.unilateralis
1048:
1045:
1010:
1007:
997:
994:
989:
988:
961:
938:
915:Attachment of
911:
867:
864:
790:Ophiocordyceps
773:
770:
752:
746:
728:
722:
701:
698:
682:Ophiocordyceps
654:
651:
637:
634:
633:
632:
627:
611:
610:
605:
600:
595:
590:
585:
580:
575:
570:
565:
560:
555:
550:
545:
540:
535:
530:
525:
520:
515:
510:
505:
500:
495:
427:Ophiocordyceps
391:
386:
353:Ophiocordyceps
326:Ophiocordyceps
321:Ophiocordyceps
306:
304:Ophiocordyceps
297:
288:
285:
205:
204:
182:
181:
175:
174:
163:
152:
151:
145:
144:
137:
135:
131:
130:
127:Ophiocordyceps
123:
119:
118:
113:
109:
108:
103:
99:
98:
93:
89:
88:
83:
79:
78:
73:
69:
68:
63:
59:
58:
45:
44:
37:
36:
28:
27:
15:
9:
6:
4:
3:
2:
3564:
3553:
3550:
3548:
3545:
3543:
3540:
3538:
3535:
3533:
3530:
3528:
3525:
3523:
3520:
3518:
3517:Fungi of Asia
3515:
3513:
3510:
3508:
3505:
3504:
3502:
3485:
3480:
3476:
3472:
3467:
3463:
3458:
3452:
3448:
3447:
3445:
3443:
3439:
3431:
3425:
3421:
3416:
3412:
3408:
3403:
3399:
3395:
3390:
3386:
3382:
3377:
3373:
3369:
3364:
3360:
3356:
3351:
3347:
3343:
3338:
3334:
3330:
3325:
3321:
3317:
3312:
3308:
3304:
3298:
3293:
3287:
3283:
3278:
3272:
3268:
3267:
3265:
3263:
3259:
3255:
3250:
3244:
3241:
3238:
3234:
3231:
3230:
3219:
3213:
3209:
3204:
3202:
3198:
3194:
3189:
3184:
3179:
3174:
3170:
3166:
3162:
3158:
3154:
3149:
3145:
3141:
3136:
3131:
3127:
3123:
3119:
3115:
3111:
3106:
3102:
3098:
3093:
3088:
3083:
3078:
3074:
3070:
3066:
3061:
3060:
3047:
3043:
3036:
3022:
3018:
3011:
3003:
2999:
2994:
2989:
2984:
2979:
2975:
2971:
2968:(5): e36352.
2967:
2963:
2959:
2952:
2950:
2948:
2946:
2944:
2928:
2922:
2914:
2910:
2905:
2900:
2896:
2892:
2888:
2884:
2880:
2876:
2872:
2865:
2857:
2853:
2849:
2842:
2834:
2830:
2826:
2822:
2818:
2814:
2807:
2799:
2795:
2791:
2787:
2783:
2779:
2776:(4): 135–40.
2775:
2771:
2764:
2756:
2752:
2748:
2744:
2741:(8): 680–92.
2740:
2736:
2728:
2720:
2716:
2712:
2708:
2704:
2700:
2693:
2685:
2681:
2677:
2673:
2669:
2662:
2646:
2642:
2638:
2631:
2623:
2619:
2615:
2611:
2608:(2): 123–37.
2607:
2603:
2596:
2588:
2584:
2579:
2574:
2569:
2564:
2560:
2556:
2552:
2548:
2544:
2537:
2535:
2533:
2531:
2529:
2527:
2518:
2514:
2509:
2504:
2500:
2496:
2492:
2488:
2484:
2477:
2469:
2465:
2460:
2455:
2451:
2447:
2443:
2439:
2435:
2428:
2420:
2416:
2411:
2406:
2403:(2): 166–76.
2402:
2398:
2394:
2387:
2385:
2383:
2381:
2379:
2370:
2366:
2361:
2356:
2351:
2346:
2342:
2338:
2334:
2327:
2325:
2323:
2321:
2319:
2304:
2300:
2294:
2286:
2282:
2277:
2272:
2268:
2264:
2260:
2256:
2252:
2245:
2236:
2231:
2227:
2223:
2219:
2217:
2213:
2209:
2200:
2198:
2189:
2185:
2180:
2175:
2171:
2167:
2163:
2159:
2155:
2148:
2133:
2132:Science Daily
2129:
2123:
2115:
2111:
2106:
2101:
2096:
2091:
2087:
2083:
2079:
2072:
2070:
2068:
2066:
2064:
2062:
2060:
2045:
2044:BBC Worldwide
2041:
2037:
2030:
2022:
2018:
2013:
2008:
2003:
1998:
1994:
1990:
1986:
1979:
1977:
1975:
1973:
1971:
1969:
1967:
1958:
1954:
1950:
1946:
1942:
1938:
1933:
1928:
1924:
1920:
1917:(3): 424–33.
1916:
1912:
1908:
1901:
1899:
1897:
1882:
1881:
1876:
1869:
1867:
1865:
1863:
1854:
1850:
1845:
1840:
1836:
1832:
1828:
1824:
1820:
1813:
1811:
1809:
1807:
1805:
1796:
1792:
1787:
1782:
1778:
1774:
1770:
1766:
1762:
1760:
1756:
1747:
1745:
1743:
1734:
1730:
1725:
1720:
1715:
1710:
1706:
1702:
1699:(3): e17024.
1698:
1694:
1690:
1683:
1681:
1672:
1668:
1663:
1658:
1653:
1648:
1644:
1640:
1636:
1629:
1627:
1625:
1623:
1621:
1619:
1617:
1615:
1613:
1611:
1609:
1607:
1598:
1594:
1589:
1584:
1580:
1576:
1572:
1568:
1564:
1557:
1555:
1553:
1544:
1540:
1536:
1532:
1529:(3): 217–24.
1528:
1524:
1516:
1514:
1512:
1510:
1508:
1493:
1489:
1485:
1481:
1474:
1460:
1456:
1455:
1450:
1444:
1440:
1431:
1430:
1426:
1425:
1419:
1417:
1413:
1412:
1406:
1403:
1399:
1395:
1391:
1387:
1386:
1379:
1375:
1371:
1367:
1366:
1361:
1357:
1356:
1345:
1342:
1338:
1334:
1329:
1327:
1319:
1315:
1313:
1303:
1300:
1296:
1291:
1290:and dryness.
1289:
1285:
1281:
1271:
1269:
1265:
1261:
1256:
1251:
1249:
1238:
1229:
1226:
1221:
1219:
1215:
1211:
1207:
1206:hyperparasite
1203:
1194:
1192:
1188:
1184:
1174:
1172:
1168:
1164:
1160:
1156:
1151:
1146:
1144:
1140:
1139:
1134:
1130:
1126:
1122:
1113:
1111:
1107:
1103:
1098:
1096:
1092:
1088:
1084:
1080:
1076:
1067:
1061:
1056:
1054:
1044:
1042:
1038:
1033:
1031:
1026:
1024:
1020:
1016:
1006:
1003:
993:
986:
982:
978:
974:
969:
965:
962:
959:
955:
952:
946:
942:
939:
936:
932:
928:
924:
920:
918:
913:
912:
910:
908:
903:
899:
893:
889:
887:
883:
877:
872:
863:
860:
857:
853:
849:
844:
842:
838:
834:
830:
826:
821:
819:
808:
806:
802:
797:
795:
791:
787:
783:
779:
769:
766:
762:
758:
750:
745:
743:
738:
734:
726:
721:
719:
715:
711:
706:
697:
695:
694:
689:
688:
683:
679:
675:
671:
666:
664:
660:
647:
642:
631:
628:
626:
623:
622:
621:
619:
617:
609:
606:
604:
601:
599:
596:
594:
591:
589:
586:
584:
581:
579:
576:
574:
571:
569:
566:
564:
561:
559:
556:
554:
551:
549:
546:
544:
541:
539:
536:
534:
531:
529:
526:
524:
521:
519:
516:
514:
511:
509:
506:
504:
501:
499:
496:
494:
491:
490:
489:
487:
485:
479:
477:
473:
469:
464:
460:
454:
452:
448:
444:
440:
435:
430:
428:
424:
420:
416:
410:
408:
404:
400:
399:
390:
385:
383:
379:
375:
371:
370:
369:Tolypocladium
365:
360:
358:
354:
350:
346:
342:
338:
334:
329:
327:
323:
322:
317:
316:
312:
305:
301:
296:
294:
284:
282:
278:
274:
269:
265:
261:
259:
255:
250:
245:
241:
238:of the tribe
237:
233:
229:
225:
221:
217:
213:
212:
203:
200:
195:
188:
183:
180:
176:
171:
167:
161:
159:
153:
150:
149:Binomial name
146:
142:
141:
136:
133:
132:
129:
128:
124:
121:
120:
117:
114:
111:
110:
107:
104:
101:
100:
97:
94:
91:
90:
87:
84:
81:
80:
77:
74:
71:
70:
67:
64:
61:
60:
55:
50:
46:
43:
38:
34:
29:
26:
22:
19:
3441:
3261:
3207:
3160:
3156:
3117:
3113:
3072:
3069:BMC Genomics
3068:
3046:The Guardian
3045:
3035:
3024:. Retrieved
3020:
3010:
2965:
2961:
2931:. Retrieved
2921:
2878:
2874:
2864:
2855:
2851:
2841:
2816:
2812:
2806:
2773:
2769:
2763:
2738:
2734:
2727:
2702:
2698:
2692:
2675:
2671:
2667:
2661:
2651:December 24,
2649:. Retrieved
2645:the original
2640:
2630:
2605:
2601:
2595:
2553:(3): e4835.
2550:
2546:
2493:(1): 67–70.
2490:
2486:
2476:
2441:
2437:
2427:
2400:
2396:
2343:(18): 4589.
2340:
2336:
2306:. Retrieved
2302:
2293:
2258:
2254:
2244:
2225:
2221:
2215:
2211:
2207:
2164:(1): 67–70.
2161:
2157:
2147:
2136:. Retrieved
2122:
2085:
2081:
2047:. Retrieved
2040:Planet Earth
2039:
2029:
1992:
1988:
1914:
1910:
1884:. Retrieved
1880:The Guardian
1878:
1826:
1822:
1768:
1764:
1758:
1754:
1696:
1692:
1642:
1638:
1570:
1566:
1526:
1522:
1495:. Retrieved
1483:
1473:
1462:. Retrieved
1452:
1443:
1427:
1415:
1410:
1407:
1401:
1393:
1383:
1369:
1368:on HBO Max,
1363:
1359:
1353:
1351:
1340:
1339:is present,
1336:
1332:
1330:
1325:
1317:
1316:
1311:
1309:
1298:
1292:
1288:UV radiation
1283:
1279:
1277:
1266:acts on its
1259:
1254:
1252:
1247:
1244:
1235:
1224:
1222:
1217:
1213:
1201:
1200:
1182:
1180:
1149:
1147:
1136:
1132:
1128:
1124:
1119:
1109:
1105:
1101:
1099:
1091:hypoglycemic
1078:
1074:
1073:
1059:
1057:
1052:
1050:
1036:
1034:
1027:
1014:
1012:
1001:
999:
991:
963:
958:hypoxanthine
948:
940:
922:
916:
914:
906:
901:
894:
890:
885:
881:
880:
875:
861:
845:
837:mitochondria
822:
809:
798:
793:
789:
786:exoskeletons
781:
777:
775:
764:
756:
754:
748:
741:
736:
732:
730:
724:
704:
703:
693:Hymenostilbe
691:
685:
681:
669:
667:
658:
656:
645:
629:
624:
615:
613:
612:
607:
602:
598:O. pulvinata
597:
592:
587:
582:
577:
572:
567:
562:
557:
552:
547:
542:
537:
532:
527:
522:
517:
512:
507:
502:
497:
492:
483:
481:
480:
475:
467:
455:
450:
446:
433:
431:
426:
414:
411:
406:
395:
393:
388:
377:
373:
367:
363:
361:
357:monophyletic
352:
348:
332:
330:
325:
319:
313:
308:
303:
299:
292:
290:
263:
262:
257:
244:pathogenesis
231:
230:ecosystems.
215:
210:
209:
208:
193:
186:
185:
157:
155:
139:
138:
126:
41:
24:
18:
3389:NatureServe
3363:iNaturalist
3286:Wikispecies
2212:Hypocreales
1989:BMC Ecology
1645:: 119–160.
1573:(1): 5–59.
1378:Craig Mazin
1341:C. leonardi
1177:Polyketides
1047:Host impact
1043:(Germany).
1019:pantropical
1000:Effects of
951:sphingosine
886:C. leonardi
742:Camponotini
476:Camponotini
472:neotropical
453:subclade."
341:Hypocreales
287:Systematics
240:Camponotini
190:Tul. (1865)
106:Hypocreales
3501:Categories
3075:(1): 620.
3026:2023-02-11
2933:2013-03-04
2308:2021-01-19
2222:Mycosphere
2138:2010-09-12
2088:(1): 166.
2049:2013-04-21
1886:2010-08-22
1497:2022-04-30
1464:2011-07-19
1436:References
1398:M.R. Carey
1348:In fiction
1041:Messel pit
964:Death grip
825:Messel Pit
772:Life cycle
765:Hirsutella
737:Hirsutella
710:subspecies
687:Hirsutella
674:teleomorph
636:Morphology
588:O. ootakii
423:speciation
398:sensu lato
331:The genus
277:polyketide
86:Ascomycota
82:Division:
3457:Q64691835
3394:2.1072941
1995:(1): 13.
1771:: 13–22.
1492:0362-4331
1210:sporangia
1191:compactin
1163:cytotoxic
1097:targets.
1087:antitumor
927:chitinase
833:sarcomere
625:O. daceti
478:species.
463:septation
364:Cordyceps
349:Cordyceps
333:Cordyceps
315:Cordyceps
300:Cordyceps
249:mandibles
134:Species:
72:Kingdom:
66:Eukaryota
3479:MycoBank
3466:Fungorum
3451:Wikidata
3376:MycoBank
3337:Fungorum
3300:BioLib:
3271:Wikidata
3197:29114054
3144:32354705
3101:26285697
3002:22567151
2962:PLOS ONE
2913:19346345
2833:16231877
2798:22937549
2790:15891934
2755:21823977
2719:16175796
2622:19075836
2587:19279680
2547:PLOS ONE
2517:20719770
2468:31315924
2419:24907198
2369:31533250
2285:33049168
2188:20719770
2114:25085339
2021:21554670
1957:31283817
1949:19627240
1853:22046474
1795:32518897
1733:21399679
1693:PLOS ONE
1671:29910522
1597:18490993
1543:22959811
1454:MycoBank
1422:See also
1159:alkaline
1138:in vitro
935:protease
801:hemocoel
678:anamorph
663:pronotum
608:O. satoi
382:stromata
234:infects
218:, is an
179:Synonyms
112:Family:
62:Domain:
3430:1599624
3355:3450915
3277:Q127353
3237:UniProt
3188:5703306
3165:Bibcode
3135:7341126
3092:4545319
2993:3342268
2970:Bibcode
2904:2687288
2883:Bibcode
2578:2652714
2555:Bibcode
2508:3030878
2459:6679347
2360:6769763
2276:7657867
2179:3030878
2105:4174324
2012:3118224
1844:3204140
1786:7274273
1724:3047535
1701:Bibcode
1662:6002356
1588:2104736
1295:somatic
1187:patulin
977:calcium
968:leucine
852:mycelia
829:atrophy
718:fitness
603:O. rami
405:within
347:. When
197:(Tul.)
122:Genus:
102:Order:
92:Class:
3484:145303
3471:145303
3427:uBio:
3420:739625
3407:268505
3381:281145
3368:465287
3342:281145
3316:711976
3303:679032
3214:
3195:
3185:
3142:
3132:
3099:
3089:
3000:
2990:
2911:
2901:
2831:
2796:
2788:
2753:
2717:
2620:
2585:
2575:
2515:
2505:
2466:
2456:
2417:
2367:
2357:
2283:
2273:
2186:
2176:
2112:
2102:
2019:
2009:
1955:
1947:
1939:
1851:
1841:
1793:
1783:
1731:
1721:
1669:
1659:
1595:
1585:
1541:
1490:
1388:, its
1100:In an
1093:, and
985:myosin
931:lipase
848:hyphae
813:
761:ascoma
459:ascoma
439:clades
311:genera
201:(1883)
172:(1931)
3329:49WSK
2794:S2CID
1953:S2CID
1937:JSTOR
1321:'
1268:genes
1063:'
1051:When
981:actin
199:Sacc.
170:Petch
76:Fungi
3547:Ants
3402:NCBI
3350:GBIF
3311:BOLD
3212:ISBN
3193:PMID
3140:PMID
3097:PMID
2998:PMID
2909:PMID
2829:PMID
2786:PMID
2751:PMID
2715:PMID
2653:2022
2618:PMID
2583:PMID
2513:PMID
2464:PMID
2415:PMID
2365:PMID
2281:PMID
2184:PMID
2110:PMID
2017:PMID
1945:PMID
1849:PMID
1791:PMID
1729:PMID
1667:PMID
1593:PMID
1539:PMID
1488:ISSN
1155:acid
979:for
933:and
839:and
755:The
731:The
690:and
684:are
345:asci
318:and
236:ants
166:Tul.
3324:CoL
3235:at
3183:PMC
3173:doi
3161:114
3130:PMC
3122:doi
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