Opiliones - Knowledge

657: 396: 988: 645: 149: 1242: 410: 126: 6470: 702: 565: 1225: 688: 6458: 670: 557:), and one lateral pair (homologous to facetted eyes of horseshoe crabs and insects). This discovery suggests that the neuroanatomy of harvestmen is more primitive than derived arachnid groups, like spiders and scorpions. It also showed that the four-eyed fossil harvestman previously discovered is most likely a member of the suborder 424: 1008:

The legs continue to twitch after they are detached because 'pacemakers' are located in the ends of the first long segment (femur) of their legs. These pacemakers send signals via the nerves to the muscles to extend the leg and then the leg relaxes between signals. While some harvestman's legs twitch

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suborder, use two mechanisms. One is bobbing, for which these particular individuals bounce their bodies. It potentially serves to confuse and deflect any identification of the exact location of their bodies. This can be a deceiving mechanism to avoid predation when they are in a large aggregation of

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in opiliones probably evolved due to natural selection, while paternal care appears to be the result of sexual selection. Depending on circumstances such as temperature, the eggs may hatch at any time after the first 20 days, up to about half a year after being laid. Harvestmen variously pass through

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Armored harvestmen in Laniatores can often use their modified morphology as weapons. Many have spines on their pedipalps, back legs, or bodies. By pinching with their chelicerae and pedipalps, they can cause harm to a potential predator. Also this has been proven to increase survival against recluse

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Some animals respond to attacks by simulating an apparent death to avoid either detection or further attacks. Arachnids such as spiders practise this mechanism when threatened or even to avoid being eaten by female spiders after mating. Thanatosis is used as a second line of defence when detected by

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Domínguez, M., Escalante, I., Carrasco-Rueda, F., Figuerola-Hernández, C.E., Marta Ayup, M., Umaña, M.N., Ramos, D., González-Zamora, A., Brizuela, C., Delgado, W., Pacheco-Esquivel, J., 2016. Losing legs and walking hard: effects of autotomy and different substrates in the locomotion of harvestmen

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Predators of harvestmen include a variety of animals, including some mammals, amphibians, and other arachnids like spiders and scorpions. Opiliones display a variety of primary and secondary defences against predation, ranging from morphological traits such as body armour to behavioral responses to

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The female lays her eggs shortly after mating to several months later. Some species build nests for this purpose. A unique feature of harvestmen is that some species practice parental care, in which the male is solely responsible for guarding the eggs resulting from multiple partners, often against

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involves direct copulation. The females store the sperm, which is aflagellate and immobile, at the tip of her ovipositor. The eggs are fertilized during oviposition. The males of some species offer a secretion (nuptial gift) from their chelicerae to the female before copulation. Sometimes, the male

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material. Such a broad range is unusual in arachnids, which are typically pure predators. Most hunting harvestmen ambush their prey, although active hunting is also found. Because their eyes cannot form images, they use their second pair of legs as antennae to explore their environment. Unlike most

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Most Opiliones, except for Cyphophthalmi, have long been thought to have a single pair of camera-type eyes in the middle of the head, oriented sideways. Eyes in Cyphophthalmi, when present, are located laterally, near the ozopores. A 305-million-year-old fossilized harvestman with two pairs of eyes

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of the family Pholcidae, (commonly referred to as "daddy long-leg spiders") most likely because of their similar appearance. Harvestmen are also referred to as "shepherd spiders" in reference to how their unusually long legs reminded observers of the ways that some European shepherds used stilts to

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Shear, W.A., Jones, T.H., Guidry, H.M., Derkarabetian, S., Richart, C.H., Minor, M., Lewis, J.J., 2014. Chemical defenses in the opilionid infraorder Insidiatores: divergence in chemical defenses between Triaenonychidae and Travunioidea and within travunioid harvestmen (Opiliones) from eastern and

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The interfamilial relationships within Opiliones are not yet fully resolved, although significant strides have been made in recent years to determine these relationships. The following list is a compilation of interfamilial relationships recovered from several recent phylogenetic studies, although

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Some harvestmen have elaborate and brightly coloured patterns or appendages which contrast with the body colouration, potentially serving as an aposematic warning to potential predators. This mechanism is thought to be commonly used during daylight, when they could be easily seen by any predators.

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Autotomised legs provide a further defence from predators because they can twitch for 60 seconds to an hour after detachment. This can also potentially serve as deflection from an attack and deceive a predator from attacking the animal. It has been shown to be successful against ants and spiders.

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Autotomy is the voluntary amputation of an appendage and is employed to escape when restrained by a predator. Eupnoi individuals, more specifically sclerosomatid harvestmen, commonly use this strategy in response to being captured. This strategy can be costly because harvestmen do not regenerate

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Particular patterns or colour markings on harvestmen's bodies can reduce detection by disrupting the animals' outlines or providing camouflage. Markings on legs can cause an interruption of the leg outline and loss of leg proportion recognition. Darker colourations and patterns function as

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guards the female after copulation, and in many species, the males defend territories. In some species, males also exhibit post-copulatory behavior in which the male specifically seeks out and shakes the female's sensory leg. This is believed to entice the female into mating a second time.

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Ballesteros, Jesús A; Santibáñez-López, Carlos E; Baker, Caitlin M; Benavides, Ligia R; Cunha, Tauana J; Gainett, Guilherme; Ontano, Andrew Z; Setton, Emily V W; Arango, Claudia P; Gavish-Regev, Efrat; Harvey, Mark S; Wheeler, Ward C; Hormiga, Gustavo; Giribet, Gonzalo; Sharma, Prashant P

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position within the Arachnida is disputed; their closest relatives may be camel spiders (Solifugae) or a larger clade comprising horseshoe crabs, Ricinulei, and Arachnopulmonata (scorpions, pseudoscorpions, and Tetrapulmonata). Although superficially similar to and often misidentified as

1395:, 410 million years ago, already show characteristics like tracheae and sexual organs, indicating that the group has lived on land since that time. Despite being similar in appearance to, and often confused with, spiders, they are probably closely related to the 5587:

Pomini, A.M., Machado, G., Pinto-da-Rocha, R., Macías-Ordóñez, R., Marsaioli, A.J., 2010. Lines of defence in the harvestman Hoplobunus mexicanus (Arachnida: Opiliones): Aposematism, stridulation, thanatosis and irritant chemicals. Biochem. Syst. Ecol. 38,

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Despite their long history, few harvestman fossils are known. This is mainly due to their delicate body structure and terrestrial habitat, making them unlikely to be found in sediments. As a consequence, most known fossils have been preserved within

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Eisner, T., Alsop, D., Meinwald, J., 1978. Secretions of Opilionids, Whip Scorpions and Pseudoscorpions, in: Arthropod Venoms, Handbook of Experimental Pharmacology / Handbuch Der Experimentellen Pharmakologie. Springer, Berlin, Heidelberg, pp.

430: 636:), grows as long as 22 mm (0.87 in). The leg span of many species is much greater than the body length and sometimes exceeds 160 mm (6.3 in) and to 340 mm (13 in) in Southeast Asia. Most species live for a year. 5808:

Machado, G., Raimundo, R.L.G., Oliveira, P.S., 2000. Daily activity schedule, gregariousness, and defensive behaviour in the Neotropical harvestman Goniosoma longipes (Opiliones: Gonyleptidae): Journal of Natural History: Vol 34, No

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of Scotland, and 305-million-year-old rocks in France. These fossils look surprisingly modern, indicating that their basic body shape developed very early on, and, at least in some taxa, has changed little since that time.

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Machado, Glauco; Carrera, Patricia C.; Pomini, Armando M.; Marsaioli, Anita J. (2005). "Chemical Defence in Harvestmen (Arachnida, Opiliones): Do Benzoquinone Secretions Deter Invertebrate and Vertebrate Predators?".

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Kury, Adriano B.; Villarreal M., Osvaldo (2015-05-01). "The prickly blade mapped: establishing homologies and a chaetotaxy for macrosetae of penis ventral plate in Gonyleptoidea (Arachnida, Opiliones, Laniatores)".

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Freezing – or the complete halt of movement – has been documented in the family Sclerosomatidae. While this can mean an increased likelihood of immediate survival, it also leads to reduced food and water intake.

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for a minute, others have been recorded to twitch up to an hour. The twitching has been hypothesised to function as an evolutionary advantage by keeping the attention of a predator while the harvestman escapes.

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Houghton, J.E., Townsend, V.R., Proud, D.N., 2011. The Ecological Significance of Leg Autotomy for Climbing Temperate Species of Harvestmen (Arachnida, Opiliones, Sclerosomatidae). Southeast. Nat. 10, 579–590.

361:), the Opiliones are a distinct order that is not closely related to spiders. They can be easily distinguished from long-legged spiders by their fused body regions and single pair of eyes in the middle of the 793:

Many species of harvestmen easily tolerate members of their own species, with aggregations of many individuals often found at protected sites near water. These aggregations may number 200 individuals in the

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Gnaspini, P., Rodrigues, G.S., 2011. Comparative study of the morphology of the gland opening area among Grassatores harvestmen (Arachnida, Opiliones, Laniatores)of Zoological Systematics and Evolutionary

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Escalante, I., Albín, A., Aisenberg, A., 2013. Lacking sensory (rather than locomotive) legs affects locomotion but not food detection in the harvestman Holmbergiana weyenberghi. Can. J. Zool. 91, 726–731.

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Hara, M.R., Gnaspini, P., 2003. Comparative study of the defensive behavior and morphology of the gland opening area among harvestmen (Arachnida, Opiliones, Gonyleptidae) under a phylogenetic perspective.

465:. The most easily discernible difference between harvestmen and spiders is that in harvestmen, the connection between the cephalothorax and abdomen is broad, so that the body appears to be a single 5401:

Gnaspini, P., Cavalheiro, A.J., 1998. Chemical and Behavioral Defensces of a Neotropical Cavernicolous Harvestman: Goniosoma spelaeum (Opiliones, Laniatores, Gonyleptidae). J. Arachnol. 26, 81–90.

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behavior is likely a strategy against climatic odds, but also against predators, combining the effect of scent secretions, and reducing the probability of any particular individual being eaten.

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other arachnids, harvestmen do not have a sucking stomach or a filtering mechanism. Rather, they ingest small particles of their food, thus making them vulnerable to internal parasites such as

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Giribet, Gonzalo; Sharma, Prashant P.; Benavides, Ligia R.; Boyer, Sarah L.; Clouse, Ronald M.; De Bivort, Benjamin L.; Dimitrov, Dimitar; Kawauchi, Gisele Y.; Murienne, Jerome (2012-01-01).

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Mattoni, C.I., García Hernández, S., Botero-Trujillo, R., Ochoa, J.A., Ojanguren-Affilastro, A.A., Outeda-Jorge, S., Pinto-da-Rocha, R. and Yamaguti, H.Y., 2001. Perder la cola o la vida. In

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work). In England, the Opiliones are called harvestmen, not because they appear at that season, but from a superstitious belief that if one is killed there will be a bad harvest that year.

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Albín, A.; Toscano-Gadea, C. A. (2015). "Predation among armored arachnids: Bothriurus bonariensis (Scorpions, Bothriuridae) versus four species of harvestmen (Harvestmen, Gonyleptidae)".

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Jones, T.C., Akoury, T.S., Hauser, C.K., Moore, D., 2011. Evidence of circadian rhythm in antipredator behaviour in the orb-weaving spider Larinioides cornutus. Anim. Behav. 82, 549–555.

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was reported in 2014. This find suggested that the eyes in Cyphophthalmi are not homologous to the eyes of other harvestmen. Many cave-adapted species are eyeless, such as the Brazilian

365:. Spiders have a distinct abdomen that is separated from the cephalothorax by a constriction, and they have three to four pairs of eyes, usually around the margins of the cephalothorax. 5679:

Chelini, M.-C., Willemart, R.H., Hebets, E.A., 2009. Costs and benefits of freezing behaviour in the harvestman Eumesosoma roeweri (Arachnida, Opiliones). Behav. Processes 82, 153–159.

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Pereira, W., Elpino-Campos, A., Del-Claro, K., Machado, G., 2004. Behavioral repertory of the neotropical harvestman ilhaia cuspidata (opiliones, gonyleptidae). J. Arachnol. 32, 22–30.

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The Opiliones are known for having exceptionally long legs relative to their body size; however, some species are short-legged. As in all Arachnida, the body in the Opiliones has two

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Hansen, L.S., Gonzales, S.F., Toft, S., Bilde, T., 2008. Thanatosis as an adaptive male mating strategy in the nuptial gift–giving spider Pisaura mirabilis. Behav. Ecol. 19, 546–551.

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and is about 10 mm (0.39 in) long, whilst the male had a discernable penis. Whether both specimens belong to the same species is not definitely known. They have long legs,

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The oldest known harvestman, from the 410-million-year-old Devonian Rhynie chert, displayed almost all the characteristics of modern species, placing the origin of harvestmen in the

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Fernández R, Sharma PP, Tourinho AL, Giribet G. 2017 The Opiliones tree of life: shedding light on harvestmen relationships through transcriptomics. Proc. R. Soc. B 284: 20162340.

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Gainett, Guilherme; Klementz, Benjamin C.; Blaszczyk, Pola; Setton, Emily V.W.; Murayama, Gabriel P.; Willemart, Rodrigo; Gavish-Regev, Efrat; Sharma, Prashant P. (February 2024).

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English speakers may colloquially refer to species of Opiliones as "daddy longlegs" or "granddaddy longlegs", but this name is also used for two other distantly related groups of

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Holmberg, R.G., Angerilli, N.P.D., LaCasse, L.J., 1984. Overwintering Aggregations of Leiobunum paessleri in Caves and Mines (Arachnida, Opiliones). J. Arachnol. 12, 195–204.

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Segovia, J.M.G., Del-Claro, K., Willemart, R.H., 2015. Defences of a Neotropical harvestman against different levels of threat by the recluse spider. Behaviour 152, 757–773.

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Machado, G., Pomini, A.M., 2008. Chemical and behavioral defences of the neotropical harvestman Camarana flavipalpi (Arachnida: Opiliones). Biochem. Syst. Ecol. 36, 369–376.

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Cáceres, N.C.; Monteiro-Filho, E.L.A (2001). "Food Habits, Home Range and Activity of Didelphis aurita (Mammalia, Marsupialia) in a Forest Fragment of Southern Brazil".

2941: 2759: 426: 4919:"Reproductive Behavior of Chavesincola inexpectabilis (Opiliones, Gonyleptidae) with Description of a New and Independently Evolved Case of Paternal Care in Harvestmen" 2791: 2601: 4337:

Glauco Machado, Ricardo Pinto-da-Rocha & Gonzalo Giribet (2007). "What are harvestmen?". In Ricardo Pinto-da-Rocha, Glauco Machado & Gonzalo Giribet (ed.).

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has evolved at least three times independently: once in the clade Progonyleptoidellinae + Caelopyginae, once in the Gonyleptinae, and once in the Heteropachylinae.

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Macias-Ordonez, R., 1998. The mating system of Leiobunum vittatum Say 1821 (Arachnida: Opiliones: Palpatores): Resource defense polygyny in the striped harvestman.

2901: 2853: 2639: 2579: 2564: 401: 1415:, fossilized by fine-grained volcanic ash around 165 million years ago, is hardly discernible from modern-day harvestmen and has been placed in the extant family 2825: 2682: 2620: 2458: 1021:

clade that either drop and run, or drop and remain motionless. This is also seen when disturbing an aggregation of multiple individuals, where they all scatter.

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is located between the base of the fourth pair of legs and the abdomen, with one opening on each side. In more active species, spiracles are also found upon the

517: 2549: 2534: 507:) differs from most arachnids in that Opiliones can swallow chunks of solid food, not only liquids. The stomotheca is formed by extensions of the coxae of the 5842:

Eisner, T., Eisner, M., Siegler, M., 2005. Secret Weapons: Defenses of Insects, Spiders, Scorpions, and Other Many-legged Creatures. Harvard University Press.

2321:, or even earlier. A recent molecular study of Opiliones, however, dated the origin of the order at about 473 million years ago (Mya), during the Ordovician. 1146:

is apparently endangered through human action. So far, no harvestman has been included in any kind of a Red List in Argentina, so they receive no protection.

6076:"Evolutionary and biogeographical history of an ancient and global group of arachnids (Arachnida: Opiliones: Cyphophthalmi) with a new taxonomic arrangement" 2435: 1231:(pincer-like) chelicerae typical of harvestmen (200× magnification); these chelicerae are homologous to chelicerae that take the form of fangs in spiders or 549:

and some Laniatores revealed that harvestman in addition to a pair median eyes also have two sets of vestigial eyes: one median pair (homologous to those of

6506: 6173:"Polyphyly of Caddoidea, reinstatement of the family Acropsopilionidae in Dyspnoi, and a revised classification system of Palpatores (Arachnida, Opiliones)" 4293: 3022: 3003: 2992: 2914: 2867: 2519: 3431: 5624:

Cokendolpher, J.C., 1987. Observations on the defensive behavior of a Neotropical Gonyleptidae (Arachnida, Opiliones). Revue Arachnologique, 7, pp.59–63.

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Machado, G., Vital, D.M., 2001. On the Occurrence of Epizoic Cyanobacteria and Liverworts on a Neotropical Harvestman (Arachnida: Opiliones) | BIOTROPICA

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Huber, B.A., Eberhard, W.G., 1997. Courtship, copulation, and genital mechanics in Physocyclus globosus (Araneae, Pholcidae). Can. J. Zool. 75, 905–918.

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da Silva Souza, E.; Willemart, R.H. (2011). "Harvest-ironman: heavy armature and not its defensive secretions, protects a harvestman against a spider".

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use substances like naphthoquinones, methyl- and ethyl-ketones. These secretions have successfully protected the harvestmen against wandering spiders (

492:. Some such Opiliones only have this shield in the males. In some species, the two posterior abdominal segments are reduced. Some of them are divided 4048: 2788: 2047: 1269:, which are both also called a "daddy longlegs"). This is untrue on several counts. None of the known species of harvestmen have venom glands; their 5931: 5851:

Shultz, J.W., Pinto-da-Rocha, R., 2007. Morphology and functional anatomy. Harvest. Biol. Opiliones Harv. Univ. Press Camb. Mass. Lond. Engl. 14–61.

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Harvestmen are a scientifically neglected group. Description of new taxa has always been dependent on the activity of a few dedicated taxonomists.

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animal in the world but possesses fangs too short or a mouth too round and small to bite a human, rendering it harmless (the same myth applies to

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They also have no silk glands and therefore do not build webs. In some highly derived species, the first five abdominal segments are fused into a

6704: 4073: 3957: 3904: 3581: 2881: 2843: 2092: 1917: 1474: 4809: 979:) that are commonly mistaken for daddy long-legs (Opiliones) also exhibit this behavior when their webs are disturbed or even during courtship. 4033: 3985: 2470: 2020: 626:

Typical body length does not exceed 7 mm (0.28 in), and some species are smaller than 1 mm, although the largest known species,

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Guffey, C., 1998. Leg Autotomy and Its Potential Fitness Costs for Two Species of Harvestmen (Arachnida, Opiliones). J. Arachnol. 26, 296–302.

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Fleming, P.A., Muller, D., Bateman, P.W., 2007. Leave it all behind: a taxonomic perspective of autotomy in invertebrates. Biological Reviews.

1435:(1638–1712) by adopting Lister's term Opiliones for this order, known in Lister's days as "harvest spiders" or "shepherd spiders", from Latin 6743: 4874:

Machado, G.; Raimundo, R. L. G. (2001). "Parental investment and the evolution of subsocial behaviour in harvestmen (Arachnida: Opiliones)".

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species are known, some of which have vivid patterns in yellow, green, and black with varied reddish and blackish mottling and reticulation.

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Dias, B.C., Willemart, R.H., 2013. The effectiveness of post-contact defenses in a prey with no pre-contact detection. Zoology 116, 168–174.

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Humphreys, R.K., Ruxton, G.D., 2018. A review of thanatosis (death feigning) as an anti-predator behaviour. Behav. Ecol. Sociobiol. 72, 22.

3019: 1543: 1291:(Harvestmen of the World) in 1923, with descriptions of all species known to that time. Other important taxonomists in this field include: 1017:

Individuals that are able to detect potential threats can flee rapidly from attack. This is seen with multiple long-legged species in the

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showing its body structure and long legs: one pair of eyes and broadly joined body tagma differentiate it from similar-looking arachnids.

6848: 5138:"The spider Enoploctenus cyclothorax (Araneae, Ctenidae) avoids preying on the harvestman Mischonyx cuspidatus (Opiliones, Gonyleptidae)" 4028: 2898: 2011: 900:. This may have an aposematic role by mimicking the colouration of glandular emissions of two other quinone-producing species. Mimicry ( 1058:, that act as a shield against predators; this is the most effective defense they use which creates a strong and unpleasant taste. In 6376: 861:

used at each life stage. Many species have also been able to camouflage their bodies by covering with secretions and debris from the

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ants. However, these chemical irritants are not able to prevent four species of harvestmen being preyed upon by the black scorpion

6047:. Ed. John Parker and Basil Hartley (1992). Colchester, Essex: Harley Books. pp. 26 & 30. (Translation of the Latin original, 2418:

was shown in 2004 to be a harvestman, after residing for almost one hundred years in its own arachnid order, the "Kustarachnida".

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Primer caso de autotomía en escorpiones (Scorpiones: Buthidae). III Congreso Latinoamericano de Aracnología, Memorias y Resúmenes

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described about a third (2,260) of today's known species from the 1910s to the 1950s, and published the landmark systematic work

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Harvestmen are well known for being chemically protected. They exude strongly odored secretions from their scent glands, called

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that has yielded many other terrestrial arachnids. An unnamed long-legged harvestman was reported from the Early Cretaceous of

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Cáceres, N.C. (2002). "Food Habits and Seed Dispersal by the White-Eared Opossum, Didelphis albiventris, in Southern Brazil".

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their legs, and leg loss reduces locomotion, speed, climbing ability, sensory perception, food detection, and territoriality.

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is in rainforest patches in northwest Argentina which are in an area being dramatically destroyed by humans. The cave-living

1411:. The Opiliones have remained almost unchanged morphologically over a long period. Indeed, one species discovered in China, 6722: 6484:—A synoptic taxonomic arrangement of the order Opiliones, down to family-group level, including some photos of the families 1336: 2690:

dating from approximately 100 Mya. It has been suggested that this may be related to the Ortholasmatinae (Nemastomatidae).

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Secondary defences allow for harvestmen to escape and survive from a predator after direct or indirect contact, including

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Misslin, R., 2003. The defence system of fear: behaviour and neurocircuitry. Neurophysiol. Clin. Neurophysiol. 33, 55–66.

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Other harvestmen may exhibit mimicry to resemble other species' appearances. Some Gonyleptidae individuals that produce

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growing on their bodies that suggest potential benefits for camouflage against large backgrounds to avoid detection by

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on the surface to form two plates beside each other. The second pair of legs is longer than the others and function as

4476:"A Critical Appraisal of the Placement of Xiphosura (Chelicerata) with Account of Known Sources of Phylogenetic Error" 2677:

etc. Bartel et al, 2023 report "These new records bring the total number of Burmese amber laniatorean species to ten"

5948: 5230: 5422:"A new species of pseudotrogulus roewer and assignment of the genus to the hernandariinae (opiliones, gonyleptidae)" 6805: 6657: 1273:

are not hollowed fangs but grasping claws that are typically very small and not strong enough to break human skin.

1156:. It is included in the Catálogo Nacional de especies amenazadas (National catalog of threatened species) from the 6748: 2401:

in Scotland is almost 340 million years old. Its placement is rather uncertain, apart from it being a harvestman.

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chemical secretions. Some of these defences have been attributed and restricted to specific groups of harvestmen.

6670: 4301: 1109:. Four species of Opiliones are on the Brazilian national list of endangered species, all of them cave-dwelling: 306:. As of July 2024, over 6,650 species of harvestmen have been discovered worldwide, although the total number of 20: 6346:

Dunlop, Jason A. (2007). "Paleontology". In Ricardo Pinto-da-Rocha, Glauco Machado & Gonzalo Giribet (ed.).

5437: 4715: 1439:, "shepherd"; Lister characterized three species from England (although not formally describing them, being a 6730: 5478:"Epizoic cyanobacteria associated with a Neotropical harvestman (Opiliones: Sclerosomatidae) from Costa Rica" 893: 572:), showing the almost fused arrangement of abdomen and cephalothorax that distinguishes these arachnids from 5753:

Roth, V.D., Roth, B.M., 1984. review of appendotomy in spiders and other arachnids. Bull.-Br. Arachnol. Soc.

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A Novel Class of Defensive Compounds in Harvestmen: Hydroxy-γ-Lactones from the Phalangiid Egaenus convexus

4526:"Comprehensive Species Sampling and Sophisticated Algorithmic Approaches Refute the Monophyly of Arachnida" 1315: 493: 481: 454: 443: 6230:"World Checklist of Opiliones species (Arachnida). Part 1: Laniatores – Travunioidea and Triaenonychoidea" 5476:

Proud, Daniel N.; Wade, Ryan R.; Rock, Philip; Townsend, Victor R.; Chavez, Donald Jiménez (August 2012).

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Naturally, most finds are from comparatively recent times. More than 20 fossil species are known from the

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To deflect attacks and enhance escape, long-legged species – commonly known as daddy long-legs – from the

6587: 6564: 5928: 5421: 4634:"A Paleozoic Stem Group to Mite Harvestmen Revealed through Integration of Phylogenetics and Development" 4373:"A Paleozoic Stem Group to Mite Harvestmen Revealed through Integration of Phylogenetics and Development" 1332: 6592: 6462: 2391:, after it was discovered that living daddy-longlegs have the same arrangement of eyes as the fossils. 6810: 6797: 148: 6758: 6381: 4427:"Anatomically modern Carboniferous harvestmen demonstrate early cladogenesis and stasis in Opiliones" 587:) that secrete a peculiar-smelling fluid when disturbed. In some species, the fluid contains noxious 5305: 1295: 6838: 6481: 6351: 4342: 4269: 2293: 1356:

Early work on the developmental biology of Opiliones from the mid-20th century was resurrected by

4745: 3509: 3072: 24: 6784: 6493: 1093:). These secretions contain multiple volatile compounds that vary among individuals and clades. 803: 6517: 5300: 4690: 4608: 2911: 1349:

Since the 1990s, study of the biology and ecology of harvestmen has intensified, especially in

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the scent glands release naphthoquinones, chloro-naphthoquinones and aliphatic methyl ketones,

1037:

and have also been shown to be used as a second line of defense when restrained by a predator.

612: 5902: 4683:"Third species of Guasiniidae (Opiliones, Laniatores) with comments on familial relationships" 4582: 6779: 1328: 1282: 1261: 1180: 763: 600: 6709: 5530:"Mimicry: imitative depiction of discharged defensive secretion on carapace of an opilionid" 4336: 6626: 6405:"First identifiable Mesozoic harvestman (Opiliones: Dyspnoi) from Cretaceous Burmese amber" 5989: 5541: 5292: 5057: 5014: 4883: 4645: 4475: 4438: 4384: 2714: 2368: 1452: 1428: 1379: 1034: 992: 905: 656: 596: 215: 6771: 5688:

Field, L.H., Glasgow, S., 2001. The Biology of Wetas, King Crickets and Their Allies. CABI

4965: 4746:"Vestigial organs alter fossil placements in an ancient group of terrestrial chelicerates" 1216:

are being considered for listing as endangered species, but as yet receive no protection.

901: 8: 6833: 6555: 5980: 5410:

Edgar, A.L., 1971. Studies on the biology and ecology of Michigan Phalangida (Opiliones).

4784:"SENCKENBERG world of biodiversity | About us | Communications | Newsroom" 2485: 2466: 1130:

Several Opiliones in Argentina appear to be vulnerable, if not endangered. These include

826:

Primary defences help the harvestmen avoid encountering a potential predator and include

809:

Harvestmen clean their legs after eating by drawing each leg in turn through their jaws.

742: 5993: 5860: 5545: 5296: 5061: 5018: 4887: 4649: 4442: 4388: 6431: 6404: 6323: 6288: 6264: 6229: 6210: 6141: 6013: 5565: 5505: 5449: 5326: 5203: 5157: 5115: 5073: 5030: 4946: 4938: 4899: 4856: 4707: 4558: 3705: 3687: 3502: 3065: 2802: 2527: 1706: 1686: 1357: 415: 395: 143: 134: 1174:

are listed as endangered species in the United States. Both are from caves in central

1066:

use nitrogen-containing substances, terpenes, aliphatic ketones, and phenolics, while

857:

are multiple species with cryptic colouration that changes over ontogeny to match the

6766: 6613: 6436: 6355: 6328: 6310: 6269: 6251: 6202: 6194: 6133: 6097: 6092: 6075: 6005: 5908: 5557: 5497: 5441: 5370: 5318: 5226: 5195: 4987: 4815: 4765: 4663: 4588: 4563: 4545: 4505: 4497: 4456: 4402: 4371:

Garwood, Russell J.; Sharma, Prashant P.; Dunlop, Jason A.; Giribet, Gonzalo (2014).

4346: 3914: 3751: 3741: 3713: 3695: 3391: 2496: 2288: 1868: 1771: 1753: 1715: 1695: 1440: 1362: 1111: 921: 675: 545: 531: 497: 390: 6214: 6145: 5569: 5509: 5453: 5330: 5161: 5119: 5077: 5034: 4950: 4903: 4860: 4839:(Arachnida: Opiliones): from premating struggles to solicitous tactile engagement". 4711: 953:

suborders, with individuals becoming rigid with legs either retracted or stretched.

904:) occurring between Brazilian harvestmen that resemble others could be explained by 6426: 6418: 6318: 6300: 6259: 6241: 6184: 6125: 6087: 6017: 5997: 5549: 5489: 5433: 5310: 5259: 5207: 5187: 5149: 5107: 5065: 5022: 4977: 4966:"Reproductive Behavior in Harvestman (Arachnida): Mating Systems and Parental Care" 4930: 4891: 4848: 4757: 4699: 4653: 4587:, vol. 5: Harvester ant to leaf-cutting ant, Marshall Cavendish, p. 263, 4553: 4537: 4487: 4446: 4392: 3896: 3474: 2699: 1859: 1305: 1301: 1164: 1075: 628: 588: 5970: 5137: 5111: 4895: 6618: 6569: 5935: 5191: 4425:

Garwood, Russell J.; Dunlop, Jason A.; Giribet, Gonzalo; Sutton, Mark D. (2011).

4180: 3919: 3876: 3827: 3644: 3606: 2874: 2770: 2702: 2587: 2572: 2363:

is known from two specimens, one a female, the other a male. The female bears an

2298: 2235: 1877: 1830: 1663: 1532: 1503: 1416: 1342: 987: 873: 738: 616: 439: 279: 6735: 1321: 6665: 6652: 5529: 4139: 4103: 4078: 3881: 3871: 3746: 3626: 3479: 3383: 3373: 3368: 3228: 3203: 3138: 2971: 2891: 2443: 2186: 2139: 2101: 1839: 1821: 1762: 1523: 1400: 1170: 1153: 1135: 917: 787: 644: 550: 6500: 6001: 5553: 5314: 5069: 5026: 4982: 4852: 4835:

Fowler-Finn, K.D.; Triana, E.; Miller, O.G. (2014). "Mating in the harvestman

4761: 4658: 4633: 4397: 4372: 2706: 2336:, and the latter having probably diverged from the Dyspnoi more than 300 Mya. 333:

Representatives of each extant suborder can be found on all continents except

6827: 6314: 6305: 6255: 6198: 6137: 6101: 5561: 5501: 5445: 4991: 4549: 4541: 4501: 4149: 4111: 4023: 4015: 3822: 3812: 3799: 3662: 3594: 3570: 2949: 2822: 2687: 2670: 2651: 2632: 2613: 2405: 2376: 2325: 2197: 2148: 2002: 1993: 1654: 1636: 1618: 1575: 1492: 1464: 1432: 1350: 1241: 1232: 1228: 1046:

spiders by causing injury, allowing the harvestman to escape from predation.

869: 771: 767: 750: 620: 446: 362: 349: 327: 311: 252: 231: 73: 6487: 6159: 5264: 5247: 4492: 4474:

Ballesteros, Jesús A; Sharma, Prashant P (2019-11-01). Halanych, Ken (ed.).

3494: 2332:

much older than 50 million years are known, despite the former presenting a

6540: 6440: 6422: 6332: 6273: 6206: 6009: 5953: 5322: 5199: 4769: 4667: 4567: 4509: 4460: 4406: 4175: 4157: 4093: 4083: 4068: 4063: 4058: 3995: 3909: 3863: 3854: 3817: 3807: 3409: 2833: 2816: 2504: 2447: 2226: 2206: 2128: 2110: 2083: 2074: 2065: 1964: 1812: 1803: 1645: 1627: 1392: 1252: 1090: 1030: 929: 536: 341: 307: 261: 1138:, the highest peak in the Sierras Chicas chain (provincia de Cordoba) and 409: 6691: 6579: 6549: 4522: 4190: 4185: 3977: 3944: 3935: 3886: 3733: 3718: 3634: 3396: 3360: 3113: 2934: 2557: 2542: 2333: 2253: 2244: 1946: 1899: 1890: 1848: 1744: 1724: 1552: 1105:

species (of all animal taxa) are considered to be at least threatened in

1102: 933: 862: 831: 661: 580: 543:. However, recent work studying the embryonic development of the species 540: 522: 470: 462: 287: 190: 48: 6246: 4942: 4918: 1458:

the placement and even monophyly of several taxa are still in question.

310:

species may exceed 10,000. The order Opiliones includes five suborders:

6228:

Kury, Adriano B.; Mendes, Amanda Cruz; Souza, Daniele R. (2014-11-05).

4451: 4426: 4200: 4038: 4005: 4000: 3990: 3952: 3843: 3756: 3614: 3469: 3287: 3146: 3046: 3033: 2745: 2364: 2329: 2271: 2029: 1982: 1973: 1955: 1908: 1793: 1780: 1512: 1270: 1266: 1213: 1084: 975:

individuals, which are all trembling at the same time. Cellar spiders (

950: 877: 848: 795: 710: 695: 633: 592: 554: 504: 474: 373: 334: 323: 246: 125: 93: 58: 6605: 6189: 6172: 6129: 5477: 4964:

Machado, Glauco; Requena, Gustavo S.; Buzatto, Bruno A. (2009-12-06).

4934: 2412:

species are currently grouped as Dyspnoi, they look more like Eupnoi.

6696: 6639: 5973:"Harvestmen (Arachnida: Opiliones) from the Middle Jurassic of China" 5493: 5153: 4195: 4129: 4121: 4088: 4043: 3781: 3723: 3639: 3589: 3547: 3541: 3328: 3265: 3211: 3159: 3107: 2985: 2957: 2864: 2781: 2749: 2710: 2508: 2398: 2348: 2301:

is sometimes misplaced in the Phalangioidea. It is not a harvestman.

2262: 2168: 2159: 2119: 2038: 1733: 1598: 1561: 1483: 1408: 1366:

as a model system for the study of arachnid comparative genomics and

1236: 1018: 976: 853: 783: 746: 731: 722: 701: 680: 469:

structure. Other differences include the fact that Opiliones have no

377: 369: 180: 160: 98: 42: 6511: 6490:—Diagnostic photographs and information on North American harvestmen 539:) from caves, most species of Cyphophthalmi, and all species of the 6683: 6644: 6600: 6534: 6469: 4703: 4134: 3789: 3253: 3184: 3088: 3057: 2812: 2766: 2734: 2695: 2489: 2451: 2344: 2340: 2318: 2177: 1607: 1404: 1396: 1389: 1385: 1224: 925: 897: 866: 608: 564: 508: 489: 340:

Well-preserved fossils have been found in the 400-million-year-old

283: 200: 115: 111: 88: 83: 68: 63: 53: 19:

For other arthropods called "daddy longlegs", and other uses, see

3678: 3295: 2924: 2753: 2666: 2647: 2628: 2609: 2422: 1676: 1311: 1055: 946: 858: 835: 827: 749:

species in the suborder Cyphophthalmi, where the males deposit a

584: 526: 458: 450: 358: 319: 241: 103: 78: 5528:

González, Andrés; Rossini, Carmen; Eisner, Thomas (2004-03-01).

5248:"Determining the fitness consequences of antipredation behavior" 4609:"4-Eyed Daddy Longlegs Helps Explain Arachnid Evolution (Video)" 6457: 3772: 3535: 3240: 3101: 2840: 2726: 2512: 2388: 1588: 1106: 971: 799: 779:

to reach maturity, with most known species having six instars.

776: 754: 714: 687: 573: 558: 354: 315: 236: 170: 6496:—Diagnostic photographs and information on European harvestmen 4743: 4249:

Kury, A. et al. (2023). WCO-Lite: World Catalogue of Opiliones

865:

found in their environments. Some hard-bodied harvestmen have

500:

or feelers. In short-legged species, this may not be obvious.

6792: 5369:. Cambridge, MA: Harvard University Press. pp. 374–399. 5281: 3341: 3042: 2371:, and no median eyes. Together with the 305-million-year-old 2311: 1256: 1212:

from around springs and other restricted habitats of central

1175: 1157: 726: 718: 649: 604: 485: 6631: 5904:

The Earwig's Tail: a modern bestiary of multi-legged legends

5223:

Defence in Animals : a Survey of Anti-predator Defences

4298:

Joel Hallan's Biology Catalog (Archive version. August 2017)

669: 6289:"A revised dated phylogeny of the arachnid order Opiliones" 6073: 5971:

Diying Huang, Paul A. Selden & Jason A. Dunlop (2009).

5438:

10.1636/0161-8202(2002)030[0173:ansopr]2.0.co;2

5419: 3401: 3038: 2425:

are possibly harvestmen, but these are not well preserved.

1029:

Multiple species within the Laniatores and Dyspnoi possess

691: 466: 5135: 4631: 4424: 4370: 1247:

Opilionid chelae shown in context to the rest of the body.

6377:"Ancient four-eyed wonder resolves daddy longleg mystery" 945:

a potential predator and is commonly observed within the

5365:

Gnaspini, P.; Hara, M. R. (2007). "Defense mechanisms".

4632:

Garwood, RJ; Sharma, PP; Dunlop, JA; Giribet, G (2014).

6507:

Joel Hallan's Biology Catalog (Archive only before 2017

5420:

Leandro Firmo, Carlos; Pinto-da-Rocha, Ricardo (2002).

5097: 4834: 2999:(Eupnoi: Sclerosomatidae) — Florissant, USA (Oligocene) 683:(the many small segments making up the end of each leg) 381:

better observe their wandering flocks from a distance.

4244: 2686:, a long-legged Dyspnoi with large eyes, was found in 5527: 5475: 5047: 4963: 2408:

of North America and Europe. While the two described

3053: 725:, feeding upon dead organisms, bird dung, and other 2694:Currently, no fossil harvestmen are known from the 1345:, Adriano Brilhante Kury, Tone Novak (21st century) 786:and colored in hues of brown, although a number of 434:

Harvestmen (Opiliones sp.) filmed in Hesse, Germany

5732:in the genus Prionostemma. J. Arachnol. 44, 76–82. 5177: 4681:Pinto-da-Rocha, Ricardo; Kury, Adriano B. (2003). 4680: 6114: 4473: 2725:Unless otherwise noted, all species are from the 6825: 6480:Adriano Kury: National Museum of Rio de Janeiro 2266:Pérez González, Kury & Alonso-Zarazaga, 2007 6227: 5891:western North America | Journal of Arachnology. 5136:Willemart, R.H.; Pellegatti-Franco, F. (2006). 4916: 4873: 4580: 2404:From about 300 Mya, several finds are from the 1070:use alkylated phenolics and benzoquinones, and 579:Harvestmen have a pair of prosomatic defensive 488:, which in most such species is fused with the 6287:Sharma, Prashant P.; Giribet, Gonzalo (2014). 4606: 3556: 851:when they remain motionless. Within the genus 6402: 6286: 6171:Groh, Selina; Giribet, Gonzalo (2015-06-01). 5894: 5245: 3524: 3510: 3073: 5364: 2575:) — Upper Carboniferous (Commentary, France) 1286: 1240: 6403:Giribet, Gonzalo; Dunlop, Jason A. (2005). 6170: 4917:Nazareth, Taís M.; Machado, Glauco (2009). 2590:) — Upper Carboniferous (Mazon Creek, U.S.) 4574: 3517: 3503: 3080: 3066: 883: 619:, unlike other arachnids. All species lay 124: 6430: 6322: 6304: 6263: 6245: 6188: 6160:https://dx.doi.org/10.1098/rspb.2016.2340 6118:Zoological Journal of the Linnean Society 6091: 6080:Biological Journal of the Linnean Society 5907:. Harvard University Press. p. 143. 5900: 5304: 5263: 4981: 4657: 4557: 4491: 4450: 4396: 1427:The Swedish naturalist and arachnologist 896:secretions have orange markings on their 5966: 5964: 3029:(Laniatores: Samoidae) — Dominican amber 2954:Florissant Fossil Beds National Monument 2920:(Dyspnoi: Nemastomatidae) — Baltic amber 2907:(Dyspnoi: Nemastomatidae) — Baltic amber 2798:(Laniatores: Samoidae) — Dominican amber 2698:. So far, they are also absent from the 1327:James C. Cokendolpher, Raymond Forster, 1223: 986: 812: 700: 686: 668: 655: 643: 563: 422: 408: 394: 6844:Extant Early Devonian first appearances 6062:Curious Facts in the History of Insects 5946: 5901:Berenbaum, May R. (30 September 2009). 5220: 5004: 4366: 4364: 4362: 4332: 4330: 4328: 4326: 4324: 4322: 4320: 4318: 4237: 2526:(Tetrophthalmi) — Upper Carboniferous ( 1255:claims that the harvestman is the most 1152:has only been found in one cave in the 6826: 6374: 6345: 5829: 5827: 5817: 5815: 5786: 5784: 5774: 5772: 5749: 5747: 5709: 5707: 5705: 5703: 5675: 5673: 5360: 4807: 4739: 4737: 4735: 4420: 4418: 4416: 3037:species (Eupnoi: Phalangiidae) — near 2860:(Eupnoi: Phalangiidae?) — Baltic amber 2849:(Eupnoi: Phalangiidae?) — Baltic amber 611:on the ventral cephalothorax, and the 6516: 6515: 6045:Martin Lister's English Spiders, 1678 5961: 5645: 5643: 5632: 5630: 5583: 5581: 5579: 5523: 5521: 5519: 5397: 5395: 5358: 5356: 5354: 5352: 5350: 5348: 5346: 5344: 5342: 5340: 5277: 5275: 5246:Lind, Johan; Cresswell, Will (2005). 5173: 5171: 5131: 5129: 5093: 5091: 5089: 5087: 4286: 3498: 3458: 3126: 3061: 911: 372:, the crane flies of the superfamily 23:. For the novel by Jean Webster, see 6798:022087FF-3A6B-980B-4FDC-84C4FD85F944 6759:be640fe4-ffd8-4b4d-a2bf-9323c3b71ff0 6671:535a87e2-e40b-4209-9156-f7f119df3b16 6348:Harvestmen: the Biology of Opiliones 5367:Harvestmen: The Biology of Opiliones 4607:Blaszczak-Boxe, Agata (2014-04-10). 4359: 4339:Harvestmen: the Biology of Opiliones 4315: 3087: 2717:, Australia, which may be a Eupnoi. 2428: 2387:are in fact members of the suborder 1579:Karaman, Mitov & Snegovaya, 2024 1096: 717:and all kinds of plant material and 5824: 5812: 5781: 5769: 5744: 5700: 5670: 4732: 4524:(2022-02-03). Teeling, Emma (ed.). 4413: 1431:(1801–1875) honored the naturalist 1407:; these four orders form the clade 821: 766:, and cleaning the eggs regularly. 13: 6849:Taxa named by Carl Jakob Sundevall 6410:Proceedings of the Royal Society B 5640: 5627: 5576: 5516: 5392: 5337: 5272: 5168: 5126: 5084: 1368:evolutionary-developmental biology 798:, and more than 70,000 in certain 741:species do occur, most harvestmen 14: 6860: 6450: 6375:Tepper, Fabien (April 11, 2014). 3054:Further internal navigation links 1184:from a cave in central Texas and 6468: 6456: 6093:10.1111/j.1095-8312.2011.01774.x 4876:Ethology Ecology & Evolution 4584:Insects and spiders of the world 4267: 3439:are possibly paraphyletic groups 2556:(Eupnoi) — Upper Carboniferous ( 2541:(Eupnoi) — Upper Carboniferous ( 2304: 1324:, William Sørensen (1880s–1890s) 1219: 1134:, which is found only on top of 1078:), wolf spiders (Lycosidae) and 705:Gregarious behavior in Opiliones 679:, showing the long legs and the 268:5 suborders, > 6,650 species 147: 46: 6396: 6368: 6339: 6280: 6221: 6164: 6152: 6108: 6067: 6054: 6038: 5940: 5921: 5884: 5875: 5865: 5854: 5845: 5836: 5802: 5793: 5756: 5735: 5725: 5716: 5691: 5682: 5661: 5652: 5618: 5609: 5600: 5591: 5469: 5460: 5413: 5404: 5383: 5239: 5214: 5050:Stud. Neotropical Fauna Environ 5041: 5007:Stud. Neotropical Fauna Environ 4998: 4957: 4910: 4867: 4828: 4801: 4776: 4674: 4625: 4600: 4530:Molecular Biology and Evolution 2965:Proholoscotolemon nemastomoides 2796:Cokendolpher & Poinar, 1998 1834:Derkarabetian & Hedin, 2018 1337:Cândido Firmino de Mello-Leitão 1024: 477:, so pose no danger to humans. 21:Daddy longlegs (disambiguation) 5947:Perkins, Sid (June 23, 2009). 4814:. Princeton University Press. 4808:Cowles, Jillian (2018-06-12). 4516: 4467: 4261: 2347:, and at least seven from the 1852:Absolon & Kratochvil, 1932 1816:Absolon & Kratochvil, 1932 1446: 1132:Pachyloidellus fulvigranulatus 1040: 384: 1: 5949:"Long-lasting daddy longlegs" 5112:10.1016/j.anbehav.2010.09.023 4896:10.1080/08927014.2001.9522780 4581:Joyce Tavolacci, ed. (2003), 4270:"Classification of Opiliones" 4231: 3015:— Florissant, USA (Oligocene) 2297:) from underground waters in 1631:Kury & Cokendolpher, 2020 939: 278:(formerly Phalangida) are an 5192:10.1016/j.beproc.2015.10.003 3430:Classification is based on 2478:Echinopustulus samuelnelsoni 2354: 1422: 1373: 1329:Clarence and Marie Goodnight 1316:Tord Tamerlan Teodor Thorell 511:and the first pair of legs. 330:, which were named in 2014. 7: 6501:The Rhynie Chert Harvestmen 6494:Harvestman: Order Opiliones 6488:Harvestman: Order Opiliones 6482:Classification of Opiliones 5285:Journal of Chemical Ecology 3459: 3290:(shorttailed whipscorpions) 2720: 2594: 1680:Hansen & Sørensen, 1904 1592:Hansen & Sørensen, 1904 1496:Hansen & Sørensen, 1904 1333:Reginald Frederick Lawrence 1276: 1049: 1035:intraspecific communication 982: 956: 639: 617:male Opiliones have a penis 413:North European harvestman ( 10: 6865: 4923:The Journal of Arachnology 3127: 2830:(Koch & Berendt, 1854) 1450: 1377: 1033:organs, which are used as 1012: 965: 841: 388: 18: 6524: 6382:Christian Science Monitor 6234:Biodiversity Data Journal 6002:10.1007/s00114-009-0556-3 5554:10.1007/s00049-003-0252-2 5315:10.1007/s10886-005-7611-0 5070:10.1076/snfe.36.2.85.2138 5027:10.1076/snfe.37.2.97.8582 4983:10.4257/oeco.2009.1301.05 4853:10.1163/1568539x-00003209 4762:10.1016/j.cub.2024.02.011 4659:10.1016/j.cub.2014.03.039 4398:10.1016/j.cub.2014.03.039 4274:www.museunacional.ufrj.br 4166: 4148: 4120: 4102: 4014: 3976: 3943: 3932: 3895: 3862: 3851: 3840: 3798: 3780: 3769: 3732: 3704: 3686: 3675: 3653: 3623: 3603: 3578: 3567: 3530: 3525:Extant Opiliones Families 3465: 3454: 3426: 3382: 3359: 3340: 3324: 3315: 3277: 3268:(tailless whip scorpions) 3261: 3252: 3236: 3227: 3196: 3169: 3158: 3137: 3133: 3122: 3096: 2942:Petrunkevitchiana oculata 2760:Philacarus hispaniolensis 2507:) — Upper Carboniferous ( 2359:The 410-million-year-old 2210:Özdikmen & Kury, 2007 2201:Özdikmen & Kury, 2007 2024:Kury & Carvalho, 2020 1288:Die Weberknechte der Erde 996:with several missing legs 936:and chemical secretions. 713:, eating primarily small 607:of the legs. They have a 267: 260: 228: 223: 144:Scientific classification 142: 132: 123: 34: 6499:University of Aberdeen: 6352:Harvard University Press 6306:10.3389/fgene.2014.00255 5389:Cokendolpher pers. comm. 4343:Harvard University Press 2665:(Insidiatores indet.) — 2602:Burmalomanius circularis 2375:, it forms the suborder 2294:Stygophalangium karamani 1986:Sharma and Giribet, 2011 1968:Sharma and Giribet, 2011 1921:Sharma and Giribet, 2011 1121:Pachylospeleus strinatii 5927:The Spider Myths Site: 2397:from East Kirkton near 1384:Harvestmen are ancient 1190:Microcina edgewoodensis 884:Aposematism and mimicry 561:(true daddy-longlegs). 503:The feeding apparatus ( 25:Daddy-Long-Legs (novel) 6423:10.1098/rspb.2005.3063 5482:Journal of Arachnology 5426:Journal of Arachnology 4691:Journal of Arachnology 4542:10.1093/molbev/msac021 2854:Cheiromachus coriaceus 2640:Mesodibunus tourinhoae 2580:Nemastomoides longipes 2565:Nemastomoides elaveris 2421:Some fossils from the 2257:Kury & Pérez, 2002 2230:Kury & Pérez, 2003 1799:(about 4,200 species) 1594:(about 1,800 species) 1296:Pierre André Latreille 1287: 1248: 1245: 1125:Spaeleoleptes spaeleus 997: 775:four to eight nymphal 706: 698: 684: 666: 653: 595:, and breathe through 576: 435: 420: 406: 402:Pachyloidellus goliath 6780:Paleobiology Database 6293:Frontiers in Genetics 5265:10.1093/beheco/ari075 5225:. New York: Longman. 4493:10.1093/sysbio/syz011 4431:Nature Communications 3149:(microwhip scorpions) 3013:(Petrunkevitch, 1922) 2946:(Petrunkevitch, 1922) 2780:species (Laniatores: 2683:Halitherses grimaldii 2621:Petroburma tarsomeria 2539:(Petrunkevitch, 1913) 2459:Brigantibunum listoni 2395:Brigantibunum listoni 2248:Gonzalez-Sponga, 1997 1283:Carl Friedrich Roewer 1262:Pholcus phalangioides 1244: 1227: 1181:Texella cokendolpheri 1144:Picunchenops spelaeus 990: 813:Antipredator defences 704: 690: 672: 665:sp. cleaning its legs 659: 647: 568:A harvestman (a male 567: 518:Caecobunus termitarum 433: 412: 398: 6666:Fauna Europaea (new) 6465:at Wikimedia Commons 6354:. pp. 247–265. 6049:Tractatus de Araneis 5221:Edmunds, M. (1974). 4847:(12–13): 1663–1686. 3331:(hooded tickspiders) 2988:) — Bitterfeld amber 2550:Kustarachne tenuipes 2535:Kustarachne longipes 2416:Kustarachne tenuipes 1682:(about 400 species) 1571:Infraorder (indet). 1470:(about 200 species) 1453:Harvestman phylogeny 1429:Carl Jakob Sundevall 1380:Harvestman phylogeny 1140:Pachyloides borellii 993:Rilaena triangularis 906:convergent evolution 745:. Except from small 648:Harvestman eating a 399:Chilean harvestman ( 6417:(1567): 1007–1013. 6247:10.3897/BDJ.2.e4094 5994:2009NW.....96..955H 5981:Naturwissenschaften 5546:2004Checo..14....5G 5297:2005JCEco..31.2519M 5062:2001SNFE...36...85C 5019:2002SNFE...37...97C 4970:Oecologia Australis 4888:2001EtEcE..13..133M 4756:(6): 1258–1270.e5. 4650:2014CBio...24.1017G 4443:2011NatCo...2..444G 4389:2014CBio...24.1017G 2784:) — Dominican amber 2663:Bartel et al, 2023 2644:Bartel et al, 2023 2625:Bartel et al, 2023 2606:Bartel et al, 2023 2584:Petrunkevitch, 1913 2486:Upper Carboniferous 2467:Early Carboniferous 2436:Eophalangium sheari 2361:Eophalangium sheari 1388:. Fossils from the 1080:Formica exsectoides 591:. They do not have 5934:2007-07-14 at the 5252:Behavioral Ecology 4837:Lieobunum vittatum 4480:Systematic Biology 4452:10.1038/ncomms1458 3706:Ischyropsalidoidea 3688:Acropsopilionoidea 3204:Pseudoscorpionida 3027:Cokendolpher, 1987 3004:Amauropilio lacoei 2993:Amauropilio atavus 2528:Montceau-les-Mines 2520:Hastocularis argus 2442:(Tetrophthalmi) — 2373:Hastocularis argus 2324:No fossils of the 1707:Ischyropsalidoidea 1687:Acropsopilionoidea 1413:Mesobunus martensi 1360:, who established 1358:Prashant P. Sharma 1335:, Jochen Martens, 1249: 1246: 1210:Microcina tiburona 998: 912:Secondary defences 764:egg-eating females 743:reproduce sexually 707: 699: 685: 667: 654: 577: 484:shield called the 436: 421: 416:Leiobunum rotundum 407: 135:Hadrobunus grandis 16:Order of arachnids 6821: 6820: 6767:Open Tree of Life 6518:Taxon identifiers 6461:Media related to 6361:978-0-674-02343-7 6190:10.1111/cla.12087 6130:10.1111/zoj.12225 5914:978-0-674-05356-4 5376:978-0-674-02343-7 5291:(11): 2519–2539. 4935:10.1636/ST08-32.1 4821:978-1-4008-9018-7 4811:Amazing Arachnids 4594:978-0-7614-7334-3 4352:978-0-674-02343-7 4345:. pp. 1–13. 4268:Kury, Adriano B. 4225: 4224: 4221: 4220: 4217: 4216: 4213: 4212: 4209: 4208: 3928: 3927: 3915:Synthetonychiidae 3836: 3835: 3765: 3764: 3752:Nipponopsalididae 3742:Dicranolasmatidae 3714:Ischyropsalididae 3696:Acropsopilionidae 3671: 3670: 3492: 3491: 3488: 3487: 3450: 3449: 3446: 3445: 3422: 3421: 3418: 3417: 3405: 3392:Opilioacariformes 3352: 3332: 3311: 3310: 3307: 3306: 3299: 3291: 3269: 3244: 3223: 3222: 3215: 3207: 3206:(pseudoscorpions) 3188: 3180: 3150: 3028: 3014: 2998: 2997:(Cockerell, 1907) 2983: 2979:Siro platypedibus 2969: 2947: 2932: 2919: 2906: 2889: 2872: 2859: 2848: 2831: 2810: 2797: 2764: 2743: 2664: 2645: 2627:(Petrobunidae) — 2626: 2607: 2585: 2570: 2555: 2540: 2525: 2502: 2497:Eotrogulus fayoli 2483: 2464: 2441: 2429:Described species 2343:, three from the 2289:Stygophalangiidae 2276: 2267: 2258: 2249: 2240: 2231: 2222: 2211: 2202: 2191: 2182: 2173: 2164: 2153: 2144: 2133: 2124: 2115: 2106: 2097: 2088: 2079: 2070: 2061: 2052: 2043: 2034: 2025: 2016: 2007: 1998: 1987: 1978: 1969: 1960: 1951: 1940: 1931: 1922: 1913: 1904: 1895: 1882: 1873: 1869:Synthetonychiidae 1864: 1853: 1844: 1835: 1826: 1817: 1808: 1798: 1785: 1776: 1772:Nipponopsalididae 1767: 1758: 1754:Dicranolasmatidae 1749: 1738: 1729: 1720: 1716:Ischyropsalididae 1711: 1700: 1696:Acropsopilionidae 1691: 1681: 1668: 1659: 1650: 1641: 1632: 1623: 1612: 1603: 1593: 1580: 1566: 1557: 1548: 1537: 1528: 1517: 1508: 1497: 1488: 1479: 1469: 1363:Phalangium opilio 1331:, Jürgen Gruber, 1112:Giupponia chagasi 1097:Endangered status 902:Müllerian mimicry 782:Most species are 709:Many species are 676:Phalangium opilio 570:Phalangium opilio 546:Phalangium opilio 532:Giupponia chagasi 431: 391:Opiliones anatomy 272: 271: 219: 118: 6856: 6814: 6813: 6801: 6800: 6788: 6787: 6775: 6774: 6762: 6761: 6752: 6751: 6739: 6738: 6736:NBNSYS0000160574 6726: 6725: 6713: 6712: 6700: 6699: 6687: 6686: 6674: 6673: 6661: 6660: 6648: 6647: 6635: 6634: 6622: 6621: 6609: 6608: 6596: 6595: 6583: 6582: 6573: 6572: 6560: 6559: 6558: 6545: 6544: 6543: 6513: 6512: 6473:Data related to 6472: 6460: 6445: 6444: 6434: 6400: 6394: 6393: 6391: 6389: 6372: 6366: 6365: 6343: 6337: 6336: 6326: 6308: 6284: 6278: 6277: 6267: 6249: 6225: 6219: 6218: 6192: 6168: 6162: 6156: 6150: 6149: 6112: 6106: 6105: 6095: 6071: 6065: 6058: 6052: 6042: 6036: 6035: 6033: 6032: 6026: 6020:. Archived from 5977: 5968: 5959: 5958: 5944: 5938: 5929:"Daddy-Longlegs" 5925: 5919: 5918: 5898: 5892: 5888: 5882: 5879: 5873: 5869: 5863: 5858: 5852: 5849: 5843: 5840: 5834: 5831: 5822: 5819: 5810: 5806: 5800: 5797: 5791: 5788: 5779: 5776: 5767: 5760: 5754: 5751: 5742: 5739: 5733: 5729: 5723: 5720: 5714: 5711: 5698: 5695: 5689: 5686: 5680: 5677: 5668: 5665: 5659: 5656: 5650: 5647: 5638: 5634: 5625: 5622: 5616: 5613: 5607: 5604: 5598: 5595: 5589: 5585: 5574: 5573: 5525: 5514: 5513: 5494:10.1636/b11-24.1 5473: 5467: 5464: 5458: 5457: 5417: 5411: 5408: 5402: 5399: 5390: 5387: 5381: 5380: 5362: 5335: 5334: 5308: 5279: 5270: 5269: 5267: 5243: 5237: 5236: 5218: 5212: 5211: 5180:Behav. Processes 5175: 5166: 5165: 5154:10.1636/S05-70.1 5133: 5124: 5123: 5095: 5082: 5081: 5045: 5039: 5038: 5002: 4996: 4995: 4985: 4961: 4955: 4954: 4914: 4908: 4907: 4871: 4865: 4864: 4832: 4826: 4825: 4805: 4799: 4798: 4796: 4795: 4786:. Archived from 4780: 4774: 4773: 4741: 4730: 4729: 4727: 4726: 4720: 4714:. Archived from 4687: 4678: 4672: 4671: 4661: 4629: 4623: 4622: 4620: 4619: 4604: 4598: 4597: 4578: 4572: 4571: 4561: 4520: 4514: 4513: 4495: 4471: 4465: 4464: 4454: 4422: 4411: 4410: 4400: 4383:(9): 1017–1023. 4368: 4357: 4356: 4334: 4313: 4312: 4310: 4309: 4300:. Archived from 4290: 4284: 4283: 4281: 4280: 4265: 4259: 4258: 4256: 4255: 4241: 3941: 3940: 3897:Triaenonychoidea 3860: 3859: 3849: 3848: 3778: 3777: 3684: 3683: 3576: 3575: 3565: 3564: 3554: 3553: 3519: 3512: 3505: 3496: 3495: 3475:Phalangiotarbida 3456: 3455: 3438: 3403: 3357: 3356: 3351: 3350: 3346: 3344: 3330: 3322: 3321: 3318: 3297: 3289: 3267: 3259: 3258: 3242: 3234: 3233: 3213: 3205: 3186: 3178: 3167: 3166: 3162: 3148: 3135: 3134: 3124: 3123: 3082: 3075: 3068: 3059: 3058: 3026: 3012: 2996: 2984:(Cyphophthalmi: 2982: 2974:) — Baltic amber 2968: 2945: 2937:) — Baltic amber 2931: 2918: 2905: 2894:) — Baltic amber 2888: 2877:) — Baltic amber 2871: 2857: 2847: 2836:) — Baltic amber 2829: 2809: 2795: 2763: 2741: 2700:Lower Cretaceous 2662: 2643: 2624: 2605: 2583: 2568: 2553: 2538: 2523: 2500: 2481: 2462: 2439: 2274: 2265: 2256: 2247: 2238: 2229: 2220: 2209: 2200: 2189: 2180: 2171: 2162: 2151: 2142: 2131: 2122: 2113: 2104: 2095: 2086: 2077: 2068: 2060:Bragagnolo, 2015 2059: 2050: 2041: 2032: 2023: 2014: 2005: 1996: 1985: 1976: 1967: 1958: 1949: 1938: 1929: 1920: 1911: 1902: 1893: 1880: 1871: 1862: 1860:Triaenonychoidea 1851: 1842: 1833: 1824: 1815: 1806: 1796: 1783: 1774: 1765: 1756: 1747: 1737:Schönhofer, 2013 1736: 1727: 1718: 1709: 1698: 1689: 1679: 1666: 1657: 1648: 1639: 1630: 1621: 1610: 1601: 1591: 1578: 1564: 1555: 1546: 1535: 1526: 1515: 1506: 1495: 1486: 1477: 1467: 1306:Maximilian Perty 1302:Carl Ludwig Koch 1290: 1165:Texella reddelli 1150:Maiorerus randoi 822:Primary defences 629:Trogulus torosus 473:glands in their 432: 214: 152: 151: 128: 109: 108: 45: 38:Temporal range: 32: 31: 6864: 6863: 6859: 6858: 6857: 6855: 6854: 6853: 6839:Arachnid orders 6824: 6823: 6822: 6817: 6809: 6804: 6796: 6791: 6783: 6778: 6770: 6765: 6757: 6755: 6747: 6742: 6734: 6729: 6721: 6716: 6708: 6703: 6695: 6690: 6682: 6677: 6669: 6664: 6656: 6651: 6643: 6638: 6630: 6625: 6617: 6612: 6604: 6599: 6591: 6586: 6578: 6576: 6568: 6563: 6554: 6553: 6548: 6539: 6538: 6533: 6520: 6453: 6448: 6401: 6397: 6387: 6385: 6373: 6369: 6362: 6344: 6340: 6285: 6281: 6226: 6222: 6169: 6165: 6157: 6153: 6113: 6109: 6072: 6068: 6059: 6055: 6043: 6039: 6030: 6028: 6024: 5975: 5969: 5962: 5945: 5941: 5936:Wayback Machine 5926: 5922: 5915: 5899: 5895: 5889: 5885: 5880: 5876: 5870: 5866: 5859: 5855: 5850: 5846: 5841: 5837: 5832: 5825: 5820: 5813: 5807: 5803: 5798: 5794: 5789: 5782: 5777: 5770: 5761: 5757: 5752: 5745: 5740: 5736: 5730: 5726: 5721: 5717: 5712: 5701: 5696: 5692: 5687: 5683: 5678: 5671: 5666: 5662: 5657: 5653: 5648: 5641: 5635: 5628: 5623: 5619: 5614: 5610: 5605: 5601: 5596: 5592: 5586: 5577: 5526: 5517: 5474: 5470: 5465: 5461: 5418: 5414: 5409: 5405: 5400: 5393: 5388: 5384: 5377: 5363: 5338: 5306:10.1.1.384.1362 5280: 5273: 5244: 5240: 5233: 5219: 5215: 5176: 5169: 5134: 5127: 5096: 5085: 5046: 5042: 5003: 4999: 4962: 4958: 4915: 4911: 4872: 4868: 4833: 4829: 4822: 4806: 4802: 4793: 4791: 4782: 4781: 4777: 4750:Current Biology 4742: 4733: 4724: 4722: 4718: 4685: 4679: 4675: 4630: 4626: 4617: 4615: 4613:livescience.com 4605: 4601: 4595: 4579: 4575: 4521: 4517: 4472: 4468: 4423: 4414: 4377:Current Biology 4369: 4360: 4353: 4335: 4316: 4307: 4305: 4292: 4291: 4287: 4278: 4276: 4266: 4262: 4253: 4251: 4243: 4242: 4238: 4234: 4228: 4226: 4205: 4181:Fissiphalliidae 4162: 4144: 4116: 4098: 4049:Cryptogeobiidae 4010: 3972: 3934: 3924: 3920:Triaenonychidae 3891: 3877:Cryptomastridae 3853: 3842: 3832: 3828:Sclerosomatidae 3794: 3771: 3761: 3728: 3700: 3677: 3667: 3655: 3649: 3645:Troglosironidae 3625: 3619: 3607:Scopulophthalmi 3605: 3599: 3580: 3569: 3558: 3552: 3526: 3523: 3493: 3484: 3461: 3442: 3436: 3414: 3378: 3348: 3347: 3345: 3342: 3336: 3316: 3303: 3273: 3248: 3219: 3214:(camel spiders) 3192: 3160: 3154: 3129: 3118: 3092: 3086: 3056: 2875:Sclerosomatidae 2789:Hummelinckiolus 2771:Dominican amber 2723: 2703:Crato Formation 2646:(Epedanidae) — 2608:(Podoctidae) — 2597: 2588:Nemastomoididae 2573:Nemastomoididae 2431: 2357: 2307: 2299:North Macedonia 2285: 2236:Fissiphalliidae 2069:Sundevall, 1833 2048:Cryptogeobiidae 1997:Sundevall, 1833 1878:Triaenonychidae 1831:Cryptomastridae 1784:Sundevall, 1833 1748:Sundevall, 1833 1664:Sclerosomatidae 1649:Latreille, 1802 1622:Latreille, 1802 1533:Troglosironidae 1504:Scopulophthalmi 1455: 1449: 1425: 1417:Sclerosomatidae 1401:pseudoscorpions 1382: 1376: 1343:Gonzalo Giribet 1279: 1222: 1198:Microcina jungi 1099: 1052: 1043: 1027: 1015: 985: 968: 959: 942: 914: 886: 844: 824: 815: 739:parthenogenetic 694:parasitising a 642: 551:horseshoe crabs 423: 393: 387: 300:harvest spiders 213: 146: 119: 107: 106: 101: 96: 91: 86: 81: 76: 71: 66: 61: 56: 51: 40: 39: 36: 28: 17: 12: 11: 5: 6862: 6852: 6851: 6846: 6841: 6836: 6819: 6818: 6816: 6815: 6802: 6789: 6776: 6763: 6753: 6740: 6727: 6714: 6701: 6688: 6675: 6662: 6653:Fauna Europaea 6649: 6636: 6623: 6610: 6597: 6584: 6574: 6561: 6546: 6530: 6528: 6522: 6521: 6510: 6509: 6504: 6497: 6491: 6485: 6478: 6477:at Wikispecies 6466: 6452: 6451:External links 6449: 6447: 6446: 6395: 6367: 6360: 6338: 6279: 6220: 6183:(3): 277–290. 6163: 6151: 6107: 6066: 6053: 6037: 5988:(8): 955–962. 5960: 5939: 5920: 5913: 5893: 5883: 5874: 5864: 5853: 5844: 5835: 5823: 5811: 5801: 5792: 5780: 5768: 5755: 5743: 5734: 5724: 5715: 5699: 5690: 5681: 5669: 5660: 5651: 5639: 5626: 5617: 5608: 5599: 5590: 5575: 5515: 5488:(2): 259–261. 5468: 5459: 5412: 5403: 5391: 5382: 5375: 5336: 5271: 5258:(5): 945–956. 5238: 5231: 5213: 5167: 5148:(3): 649–652. 5125: 5083: 5040: 4997: 4956: 4929:(2): 127–134. 4909: 4882:(2): 133–150. 4866: 4827: 4820: 4800: 4775: 4731: 4704:10.1636/H02-59 4698:(3): 394–399. 4673: 4644:(9): 1017–23. 4624: 4599: 4593: 4573: 4515: 4486:(6): 896–917. 4466: 4412: 4358: 4351: 4314: 4285: 4260: 4235: 4233: 4230: 4223: 4222: 4219: 4218: 4215: 4214: 4211: 4210: 4207: 4206: 4204: 4203: 4198: 4193: 4188: 4183: 4178: 4172: 4170: 4164: 4163: 4161: 4160: 4154: 4152: 4146: 4145: 4143: 4142: 4140:Stygnommatidae 4137: 4132: 4126: 4124: 4118: 4117: 4115: 4114: 4108: 4106: 4104:Phalangodoidea 4100: 4099: 4097: 4096: 4091: 4086: 4081: 4079:Otilioleptidae 4076: 4071: 4066: 4061: 4056: 4051: 4046: 4041: 4036: 4031: 4026: 4020: 4018: 4012: 4011: 4009: 4008: 4003: 3998: 3993: 3988: 3982: 3980: 3974: 3973: 3971: 3970: 3968:Trionyxellidae 3965: 3960: 3955: 3949: 3947: 3938: 3930: 3929: 3926: 3925: 3923: 3922: 3917: 3912: 3907: 3901: 3899: 3893: 3892: 3890: 3889: 3884: 3882:Paranonychidae 3879: 3874: 3872:Cladonychiidae 3868: 3866: 3857: 3846: 3838: 3837: 3834: 3833: 3831: 3830: 3825: 3820: 3815: 3810: 3804: 3802: 3796: 3795: 3793: 3792: 3786: 3784: 3775: 3767: 3766: 3763: 3762: 3760: 3759: 3754: 3749: 3747:Nemastomatidae 3744: 3738: 3736: 3730: 3729: 3727: 3726: 3721: 3716: 3710: 3708: 3702: 3701: 3699: 3698: 3692: 3690: 3681: 3673: 3672: 3669: 3668: 3666: 3665: 3659: 3657: 3651: 3650: 3648: 3647: 3642: 3637: 3631: 3629: 3627:Sternophthalmi 3621: 3620: 3618: 3617: 3611: 3609: 3601: 3600: 3598: 3597: 3592: 3586: 3584: 3573: 3562: 3551: 3550: 3544: 3538: 3531: 3528: 3527: 3522: 3521: 3514: 3507: 3499: 3490: 3489: 3486: 3485: 3483: 3482: 3480:Trigonotarbida 3477: 3472: 3466: 3463: 3462: 3452: 3451: 3448: 3447: 3444: 3443: 3441: 3440: 3437:Items in green 3434: 3427: 3424: 3423: 3420: 3419: 3416: 3415: 3413: 3412: 3407: 3399: 3394: 3388: 3386: 3384:Parasitiformes 3380: 3379: 3377: 3376: 3374:Trombidiformes 3371: 3369:Sarcoptiformes 3365: 3363: 3354: 3349:(ticks, mites) 3338: 3337: 3335: 3334: 3325: 3319: 3313: 3312: 3309: 3308: 3305: 3304: 3302: 3301: 3293: 3284: 3282: 3275: 3274: 3272: 3271: 3262: 3256: 3250: 3249: 3247: 3246: 3237: 3231: 3229:Tetrapulmonata 3225: 3224: 3221: 3220: 3218: 3217: 3209: 3200: 3198: 3194: 3193: 3191: 3190: 3182: 3173: 3171: 3164: 3156: 3155: 3153: 3152: 3143: 3141: 3139:Incertae sedis 3131: 3130: 3120: 3119: 3117: 3116: 3110: 3104: 3097: 3094: 3093: 3085: 3084: 3077: 3070: 3062: 3055: 3052: 3051: 3050: 3030: 3016: 3000: 2989: 2975: 2972:Cladonychiidae 2961: 2938: 2921: 2908: 2895: 2892:Nemastomatidae 2878: 2861: 2850: 2837: 2819: 2799: 2785: 2773: 2756: 2722: 2719: 2692: 2691: 2675: 2674: 2655: 2636: 2617: 2596: 2593: 2592: 2591: 2576: 2569:Thevenin, 1901 2561: 2546: 2531: 2516: 2501:Thevenin, 1901 2493: 2474: 2455: 2444:Early Devonian 2430: 2427: 2356: 2353: 2306: 2303: 2291:(one species, 2284: 2283: 2282: 2281: 2280: 2279: 2278: 2277: 2275:Sørensen, 1886 2268: 2259: 2250: 2241: 2232: 2221:Sørensen, 1886 2214: 2213: 2212: 2194: 2193: 2192: 2187:Stygnommatidae 2183: 2181:Sørensen, 1886 2174: 2163:Sørensen, 1886 2156: 2155: 2154: 2140:Phalangodoidea 2136: 2135: 2134: 2132:Sørensen, 1932 2125: 2116: 2107: 2102:Otilioleptidae 2098: 2089: 2080: 2071: 2062: 2053: 2044: 2035: 2026: 2017: 2008: 1990: 1989: 1988: 1979: 1970: 1961: 1959:Sørensen, 1886 1950:Sørensen, 1886 1943: 1942: 1941: 1936:Trionyxellidae 1932: 1923: 1914: 1912:Sørensen, 1884 1903:Sørensen, 1884 1887: 1886: 1885: 1884: 1883: 1881:Sørensen, 1886 1874: 1863:Sørensen, 1886 1856: 1855: 1854: 1845: 1840:Paranonychidae 1836: 1827: 1822:Cladonychiidae 1790: 1789: 1788: 1787: 1786: 1777: 1768: 1763:Nemastomatidae 1759: 1741: 1740: 1739: 1730: 1721: 1703: 1702: 1701: 1673: 1672: 1671: 1670: 1669: 1660: 1651: 1642: 1640:Lawrence, 1931 1633: 1615: 1614: 1613: 1585: 1584: 1583: 1582: 1581: 1569: 1568: 1567: 1558: 1540: 1539: 1538: 1524:Sternophthalmi 1520: 1519: 1518: 1500: 1499: 1498: 1489: 1460: 1451:Main article: 1448: 1445: 1424: 1421: 1378:Main article: 1375: 1372: 1347: 1346: 1340: 1339:(20th century) 1325: 1319: 1309: 1299: 1298:(18th century) 1278: 1275: 1221: 1218: 1206:Microcina lumi 1202:Microcina leei 1194:Microcina homi 1186:Calicina minor 1171:Texella reyesi 1154:Canary Islands 1136:Cerro Uritorco 1117:Iandumoema uai 1098: 1095: 1051: 1048: 1042: 1039: 1026: 1023: 1014: 1011: 984: 981: 967: 964: 958: 955: 941: 938: 913: 910: 885: 882: 843: 840: 823: 820: 814: 811: 641: 638: 389:Main article: 386: 383: 378:cellar spiders 304:daddy longlegs 270: 269: 265: 264: 258: 257: 256: 255: 249: 244: 239: 234: 226: 225: 221: 220: 208: 204: 203: 198: 194: 193: 188: 184: 183: 178: 174: 173: 168: 164: 163: 158: 154: 153: 140: 139: 130: 129: 121: 120: 102: 97: 92: 87: 82: 77: 72: 67: 62: 57: 52: 47: 37: 15: 9: 6: 4: 3: 2: 6861: 6850: 6847: 6845: 6842: 6840: 6837: 6835: 6832: 6831: 6829: 6812: 6807: 6803: 6799: 6794: 6790: 6786: 6781: 6777: 6773: 6768: 6764: 6760: 6754: 6750: 6745: 6741: 6737: 6732: 6728: 6724: 6719: 6715: 6711: 6706: 6702: 6698: 6693: 6689: 6685: 6680: 6676: 6672: 6667: 6663: 6659: 6654: 6650: 6646: 6641: 6637: 6633: 6628: 6624: 6620: 6615: 6611: 6607: 6602: 6598: 6594: 6589: 6585: 6581: 6575: 6571: 6566: 6562: 6557: 6551: 6547: 6542: 6536: 6532: 6531: 6529: 6527: 6523: 6519: 6514: 6508: 6505: 6502: 6498: 6495: 6492: 6489: 6486: 6483: 6479: 6476: 6471: 6467: 6464: 6459: 6455: 6454: 6442: 6438: 6433: 6428: 6424: 6420: 6416: 6412: 6411: 6406: 6399: 6384: 6383: 6378: 6371: 6363: 6357: 6353: 6349: 6342: 6334: 6330: 6325: 6320: 6316: 6312: 6307: 6302: 6298: 6294: 6290: 6283: 6275: 6271: 6266: 6261: 6257: 6253: 6248: 6243: 6239: 6235: 6231: 6224: 6216: 6212: 6208: 6204: 6200: 6196: 6191: 6186: 6182: 6178: 6174: 6167: 6161: 6155: 6147: 6143: 6139: 6135: 6131: 6127: 6123: 6119: 6111: 6103: 6099: 6094: 6089: 6086:(1): 92–130. 6085: 6081: 6077: 6070: 6063: 6060:Frank Cowan, 6057: 6050: 6046: 6041: 6027:on 2016-03-05 6023: 6019: 6015: 6011: 6007: 6003: 5999: 5995: 5991: 5987: 5983: 5982: 5974: 5967: 5965: 5956: 5955: 5950: 5943: 5937: 5933: 5930: 5924: 5916: 5910: 5906: 5905: 5897: 5887: 5878: 5868: 5862: 5857: 5848: 5839: 5830: 5828: 5818: 5816: 5805: 5796: 5787: 5785: 5775: 5773: 5765: 5759: 5750: 5748: 5738: 5728: 5719: 5710: 5708: 5706: 5704: 5694: 5685: 5676: 5674: 5664: 5655: 5646: 5644: 5633: 5631: 5621: 5612: 5603: 5594: 5584: 5582: 5580: 5571: 5567: 5563: 5559: 5555: 5551: 5547: 5543: 5539: 5535: 5531: 5524: 5522: 5520: 5511: 5507: 5503: 5499: 5495: 5491: 5487: 5483: 5479: 5472: 5463: 5455: 5451: 5447: 5443: 5439: 5435: 5431: 5427: 5423: 5416: 5407: 5398: 5396: 5386: 5378: 5372: 5368: 5361: 5359: 5357: 5355: 5353: 5351: 5349: 5347: 5345: 5343: 5341: 5332: 5328: 5324: 5320: 5316: 5312: 5307: 5302: 5298: 5294: 5290: 5286: 5278: 5276: 5266: 5261: 5257: 5253: 5249: 5242: 5234: 5232:0-582-44132-3 5228: 5224: 5217: 5209: 5205: 5201: 5197: 5193: 5189: 5185: 5181: 5174: 5172: 5163: 5159: 5155: 5151: 5147: 5143: 5139: 5132: 5130: 5121: 5117: 5113: 5109: 5105: 5101: 5094: 5092: 5090: 5088: 5079: 5075: 5071: 5067: 5063: 5059: 5055: 5051: 5044: 5036: 5032: 5028: 5024: 5020: 5016: 5013:(2): 97–104. 5012: 5008: 5001: 4993: 4989: 4984: 4979: 4975: 4971: 4967: 4960: 4952: 4948: 4944: 4940: 4936: 4932: 4928: 4924: 4920: 4913: 4905: 4901: 4897: 4893: 4889: 4885: 4881: 4877: 4870: 4862: 4858: 4854: 4850: 4846: 4842: 4838: 4831: 4823: 4817: 4813: 4812: 4804: 4790:on 2015-11-19 4789: 4785: 4779: 4771: 4767: 4763: 4759: 4755: 4751: 4747: 4740: 4738: 4736: 4721:on 2016-03-28 4717: 4713: 4709: 4705: 4701: 4697: 4693: 4692: 4684: 4677: 4669: 4665: 4660: 4655: 4651: 4647: 4643: 4639: 4635: 4628: 4614: 4610: 4603: 4596: 4590: 4586: 4585: 4577: 4569: 4565: 4560: 4555: 4551: 4547: 4543: 4539: 4535: 4531: 4527: 4519: 4511: 4507: 4503: 4499: 4494: 4489: 4485: 4481: 4477: 4470: 4462: 4458: 4453: 4448: 4444: 4440: 4436: 4432: 4428: 4421: 4419: 4417: 4408: 4404: 4399: 4394: 4390: 4386: 4382: 4378: 4374: 4367: 4365: 4363: 4354: 4348: 4344: 4340: 4333: 4331: 4329: 4327: 4325: 4323: 4321: 4319: 4304:on 2017-08-04 4303: 4299: 4295: 4289: 4275: 4271: 4264: 4250: 4246: 4240: 4236: 4229: 4202: 4199: 4197: 4194: 4192: 4189: 4187: 4184: 4182: 4179: 4177: 4174: 4173: 4171: 4169: 4165: 4159: 4156: 4155: 4153: 4151: 4150:Sandokanoidea 4147: 4141: 4138: 4136: 4133: 4131: 4128: 4127: 4125: 4123: 4119: 4113: 4112:Phalangodidae 4110: 4109: 4107: 4105: 4101: 4095: 4092: 4090: 4087: 4085: 4082: 4080: 4077: 4075: 4074:Nomoclastidae 4072: 4070: 4067: 4065: 4062: 4060: 4057: 4055: 4052: 4050: 4047: 4045: 4042: 4040: 4037: 4035: 4032: 4030: 4027: 4025: 4024:Agoristenidae 4022: 4021: 4019: 4017: 4016:Gonyleptoidea 4013: 4007: 4004: 4002: 3999: 3997: 3994: 3992: 3989: 3987: 3984: 3983: 3981: 3979: 3975: 3969: 3966: 3964: 3961: 3959: 3958:Pyramidopidae 3956: 3954: 3951: 3950: 3948: 3946: 3942: 3939: 3937: 3931: 3921: 3918: 3916: 3913: 3911: 3908: 3906: 3905:Buemarinoidae 3903: 3902: 3900: 3898: 3894: 3888: 3885: 3883: 3880: 3878: 3875: 3873: 3870: 3869: 3867: 3865: 3861: 3858: 3856: 3850: 3847: 3845: 3839: 3829: 3826: 3824: 3823:Protolophidae 3821: 3819: 3816: 3814: 3813:Neopilionidae 3811: 3809: 3806: 3805: 3803: 3801: 3800:Phalangioidea 3797: 3791: 3788: 3787: 3785: 3783: 3779: 3776: 3774: 3768: 3758: 3755: 3753: 3750: 3748: 3745: 3743: 3740: 3739: 3737: 3735: 3731: 3725: 3722: 3720: 3717: 3715: 3712: 3711: 3709: 3707: 3703: 3697: 3694: 3693: 3691: 3689: 3685: 3682: 3680: 3674: 3664: 3663:Parasironidae 3661: 3660: 3658: 3652: 3646: 3643: 3641: 3638: 3636: 3633: 3632: 3630: 3628: 3622: 3616: 3613: 3612: 3610: 3608: 3602: 3596: 3595:Stylocellidae 3593: 3591: 3588: 3587: 3585: 3583: 3582:Boreophthalmi 3577: 3574: 3572: 3571:Cyphophthalmi 3566: 3563: 3561: 3555: 3549: 3545: 3543: 3539: 3537: 3533: 3532: 3529: 3520: 3515: 3513: 3508: 3506: 3501: 3500: 3497: 3481: 3478: 3476: 3473: 3471: 3468: 3467: 3464: 3457: 3453: 3435: 3433: 3432:Shultz (2007) 3429: 3428: 3425: 3411: 3408: 3406: 3400: 3398: 3395: 3393: 3390: 3389: 3387: 3385: 3381: 3375: 3372: 3370: 3367: 3366: 3364: 3362: 3358: 3355: 3353: 3339: 3333: 3327: 3326: 3323: 3320: 3314: 3300: 3298:(vinegaroons) 3294: 3292: 3286: 3285: 3283: 3281: 3278:Thelyphonida 3276: 3270: 3264: 3263: 3260: 3257: 3255: 3251: 3245: 3239: 3238: 3235: 3232: 3230: 3226: 3216: 3210: 3208: 3202: 3201: 3199: 3195: 3189: 3183: 3181: 3175: 3174: 3172: 3168: 3165: 3163: 3157: 3151: 3145: 3144: 3142: 3140: 3136: 3132: 3125: 3121: 3115: 3111: 3109: 3105: 3103: 3099: 3098: 3095: 3090: 3083: 3078: 3076: 3071: 3069: 3064: 3063: 3060: 3048: 3044: 3040: 3036: 3035: 3031: 3025: 3024: 3021: 3017: 3010: 3006: 3005: 3001: 2995: 2994: 2990: 2987: 2981: 2980: 2976: 2973: 2970:(Laniatores: 2967: 2966: 2962: 2959: 2955: 2951: 2950:Phalangioidea 2944: 2943: 2939: 2936: 2930: 2929: 2926: 2922: 2917: 2916: 2913: 2909: 2904: 2903: 2900: 2896: 2893: 2887: 2886: 2883: 2882:Histricostoma 2879: 2876: 2870: 2869: 2866: 2862: 2856: 2855: 2851: 2846: 2845: 2842: 2838: 2835: 2828: 2827: 2824: 2823:Dicranopalpus 2820: 2818: 2814: 2808: 2807: 2804: 2800: 2794: 2793: 2790: 2786: 2783: 2779: 2778: 2774: 2772: 2768: 2765:(Laniatores: 2762: 2761: 2757: 2755: 2751: 2747: 2740: 2739: 2736: 2732: 2731: 2730: 2728: 2718: 2716: 2712: 2708: 2705:of Brazil, a 2704: 2701: 2697: 2689: 2688:Burmese amber 2685: 2684: 2680: 2679: 2678: 2672: 2671:Burmese amber 2668: 2661: 2660: 2656: 2653: 2652:Burmese amber 2649: 2642: 2641: 2637: 2634: 2633:Burmese amber 2630: 2623: 2622: 2618: 2615: 2614:Burmese amber 2611: 2604: 2603: 2599: 2598: 2589: 2582: 2581: 2577: 2574: 2567: 2566: 2562: 2559: 2554:Scudder, 1890 2552: 2551: 2547: 2544: 2537: 2536: 2532: 2529: 2524:Garwood, 2014 2522: 2521: 2517: 2514: 2510: 2506: 2499: 2498: 2494: 2491: 2487: 2484:(Dyspnoi?) — 2480: 2479: 2475: 2472: 2468: 2461: 2460: 2456: 2453: 2449: 2445: 2438: 2437: 2433: 2432: 2426: 2424: 2419: 2417: 2413: 2411: 2410:Nemastomoides 2407: 2406:Coal Measures 2402: 2400: 2396: 2392: 2390: 2386: 2382: 2378: 2377:Tetrophthalmi 2374: 2370: 2366: 2362: 2352: 2350: 2346: 2342: 2337: 2335: 2331: 2327: 2326:Cyphophthalmi 2322: 2320: 2315: 2313: 2305:Fossil record 2302: 2300: 2296: 2295: 2290: 2273: 2269: 2264: 2260: 2255: 2251: 2246: 2242: 2239:Martens, 1988 2237: 2233: 2228: 2224: 2223: 2219: 2215: 2208: 2204: 2203: 2199: 2198:Sandokanoidea 2195: 2188: 2184: 2179: 2175: 2172:Thorell, 1889 2170: 2166: 2165: 2161: 2157: 2150: 2149:Phalangodidae 2146: 2145: 2141: 2137: 2130: 2126: 2121: 2117: 2112: 2108: 2103: 2099: 2094: 2093:Nomoclastidae 2090: 2085: 2081: 2076: 2072: 2067: 2063: 2058: 2054: 2049: 2045: 2040: 2036: 2031: 2027: 2022: 2018: 2013: 2009: 2006:Šilhavý, 1973 2004: 2003:Agoristenidae 2000: 1999: 1995: 1994:Gonyleptoidea 1991: 1984: 1980: 1975: 1971: 1966: 1962: 1957: 1953: 1952: 1948: 1944: 1937: 1933: 1930:Martens, 2020 1928: 1924: 1919: 1918:Pyramidopidae 1915: 1910: 1906: 1905: 1901: 1897: 1896: 1892: 1888: 1879: 1875: 1872:Forster, 1954 1870: 1866: 1865: 1861: 1857: 1850: 1846: 1841: 1837: 1832: 1828: 1823: 1819: 1818: 1814: 1810: 1809: 1805: 1801: 1800: 1797:Thorell, 1876 1795: 1791: 1782: 1778: 1775:Martens, 1976 1773: 1769: 1764: 1760: 1755: 1751: 1750: 1746: 1742: 1735: 1731: 1726: 1722: 1717: 1713: 1712: 1708: 1704: 1697: 1693: 1692: 1688: 1684: 1683: 1678: 1674: 1665: 1661: 1656: 1655:Protolophidae 1652: 1647: 1643: 1638: 1637:Neopilionidae 1634: 1629: 1625: 1624: 1620: 1619:Phalangioidea 1616: 1609: 1605: 1604: 1600: 1596: 1595: 1590: 1586: 1577: 1576:Parasironidae 1573: 1572: 1570: 1563: 1559: 1554: 1550: 1549: 1545: 1541: 1534: 1530: 1529: 1527:Giribet, 2012 1525: 1521: 1514: 1510: 1509: 1507:Giribet, 2012 1505: 1501: 1494: 1493:Stylocellidae 1490: 1485: 1481: 1480: 1478:Giribet, 2012 1476: 1475:Boreophthalmi 1472: 1471: 1466: 1465:Cyphophthalmi 1462: 1461: 1459: 1454: 1444: 1442: 1438: 1434: 1433:Martin Lister 1430: 1420: 1418: 1414: 1410: 1406: 1402: 1398: 1394: 1391: 1387: 1381: 1371: 1369: 1365: 1364: 1359: 1354: 1352: 1351:South America 1344: 1341: 1338: 1334: 1330: 1326: 1323: 1320: 1318:(1860s–1870s) 1317: 1313: 1310: 1308:(1830s–1850s) 1307: 1303: 1300: 1297: 1294: 1293: 1292: 1289: 1284: 1274: 1272: 1268: 1264: 1263: 1258: 1254: 1243: 1238: 1234: 1230: 1226: 1220:Misconception 1217: 1215: 1211: 1207: 1203: 1199: 1195: 1191: 1187: 1183: 1182: 1177: 1173: 1172: 1167: 1166: 1161: 1159: 1155: 1151: 1147: 1145: 1141: 1137: 1133: 1128: 1126: 1122: 1118: 1114: 1113: 1108: 1104: 1094: 1092: 1088: 1086: 1081: 1077: 1073: 1069: 1065: 1061: 1060:Cyphophthalmi 1057: 1047: 1038: 1036: 1032: 1022: 1020: 1010: 1006: 1002: 995: 994: 989: 980: 978: 973: 963: 954: 952: 948: 937: 935: 931: 927: 923: 919: 909: 907: 903: 899: 895: 890: 881: 879: 875: 871: 870:cyanobacteria 868: 864: 860: 856: 855: 850: 839: 837: 833: 829: 819: 810: 807: 805: 801: 797: 791: 789: 785: 780: 778: 773: 772:Maternal care 769: 768:Paternal care 765: 759: 756: 752: 751:spermatophore 748: 744: 740: 735: 733: 728: 724: 720: 716: 712: 703: 697: 693: 689: 682: 678: 677: 671: 664: 663: 658: 651: 646: 637: 635: 631: 630: 624: 622: 618: 615:is direct as 614: 610: 606: 602: 598: 594: 590: 586: 582: 575: 571: 566: 562: 560: 556: 552: 548: 547: 542: 538: 534: 533: 528: 524: 520: 519: 512: 510: 506: 501: 499: 495: 491: 487: 483: 478: 476: 472: 468: 464: 460: 457:10-segmented 456: 452: 448: 447:cephalothorax 445: 441: 418: 417: 411: 404: 403: 397: 392: 382: 379: 375: 371: 366: 364: 363:cephalothorax 360: 356: 351: 346: 343: 342:Rhynie cherts 338: 336: 331: 329: 328:Tetrophthalmi 325: 321: 317: 313: 312:Cyphophthalmi 309: 305: 301: 297: 293: 289: 285: 281: 277: 266: 263: 259: 254: 253:Tetrophthalmi 250: 248: 245: 243: 240: 238: 235: 233: 232:Cyphophthalmi 230: 229: 227: 222: 217: 212: 209: 206: 205: 202: 199: 196: 195: 192: 189: 186: 185: 182: 179: 176: 175: 172: 169: 166: 165: 162: 159: 156: 155: 150: 145: 141: 137: 136: 131: 127: 122: 117: 113: 105: 100: 95: 90: 85: 80: 75: 70: 65: 60: 55: 50: 44: 33: 30: 26: 22: 6525: 6414: 6408: 6398: 6386:. Retrieved 6380: 6370: 6347: 6341: 6296: 6292: 6282: 6240:(2): e4094. 6237: 6233: 6223: 6180: 6176: 6166: 6154: 6121: 6117: 6110: 6083: 6079: 6069: 6061: 6056: 6048: 6044: 6040: 6029:. Retrieved 6022:the original 5985: 5979: 5954:Science News 5952: 5942: 5923: 5903: 5896: 5886: 5877: 5867: 5856: 5847: 5838: 5804: 5795: 5766:(pp. 83–84). 5763: 5758: 5737: 5727: 5718: 5693: 5684: 5663: 5654: 5620: 5611: 5602: 5593: 5537: 5534:Chemoecology 5533: 5485: 5481: 5471: 5462: 5429: 5425: 5415: 5406: 5385: 5366: 5288: 5284: 5255: 5251: 5241: 5222: 5216: 5183: 5179: 5145: 5141: 5103: 5099: 5056:(2): 85–92. 5053: 5049: 5043: 5010: 5006: 5000: 4976:(1): 58–79. 4973: 4969: 4959: 4926: 4922: 4912: 4879: 4875: 4869: 4844: 4840: 4836: 4830: 4810: 4803: 4792:. Retrieved 4788:the original 4778: 4753: 4749: 4723:. Retrieved 4716:the original 4695: 4689: 4676: 4641: 4637: 4627: 4616:. Retrieved 4612: 4602: 4583: 4576: 4533: 4529: 4518: 4483: 4479: 4469: 4434: 4430: 4380: 4376: 4338: 4306:. Retrieved 4302:the original 4297: 4288: 4277:. Retrieved 4273: 4263: 4252:. Retrieved 4248: 4239: 4227: 4176:Escadabiidae 4158:Sandokanidae 4094:Stygnopsidae 4084:Prostygnidae 4069:Metasarcidae 4064:Manaosbiidae 4059:Gonyleptidae 4034:Askawachidae 3996:Petrobunidae 3986:Beloniscidae 3910:Lomanellidae 3864:Travunioidea 3855:Insidiatores 3818:Phalangiidae 3808:Globipedidae 3559: 3410:Mesostigmata 3279: 3197:Haplocnemata 3179:(harvestmen) 3176: 3170:Stomothecata 3032: 3018: 3008: 3002: 2991: 2977: 2963: 2940: 2923: 2910: 2902:denticulatum 2897: 2885:tuberculatum 2880: 2863: 2852: 2839: 2834:Phalangiidae 2821: 2817:Baltic amber 2801: 2787: 2775: 2758: 2733: 2724: 2693: 2681: 2676: 2673:(Cenomanian) 2657: 2654:(Cenomanian) 2638: 2635:(Cenomanian) 2619: 2616:(Cenomanian) 2600: 2586:(Dyspnoi: † 2578: 2571:(Dyspnoi: † 2563: 2548: 2533: 2518: 2505:Eotrogulidae 2503:(Dyspnoi: † 2495: 2482:Dunlop, 2004 2476: 2471:East Kirkton 2465:(Eupnoi?) — 2463:Dunlop, 2005 2457: 2440:Dunlop, 2004 2434: 2420: 2415: 2414: 2409: 2403: 2394: 2393: 2384: 2380: 2372: 2360: 2358: 2338: 2323: 2316: 2308: 2292: 2286: 2227:Escadabiidae 2216:Superfamily 2207:Sandokanidae 2196:Superfamily 2190:Roewer, 1923 2158:Superfamily 2138:Superfamily 2129:Stygnopsidae 2114:Roewer, 1913 2111:Prostygnidae 2105:Acosta, 2019 2096:Roewer, 1943 2084:Metasarcidae 2078:Roewer, 1943 2075:Manaosbiidae 2066:Gonyleptidae 2042:Roewer, 1913 2021:Askawachidae 1992:Superfamily 1977:Roewer, 1912 1965:Petrobunidae 1945:Superfamily 1939:Roewer, 1912 1898:Superfamily 1858:Superfamily 1843:Briggs, 1971 1813:Travunioidea 1811:Superfamily 1804:Insidiatores 1743:Superfamily 1728:Dresco, 1970 1705:Superfamily 1699:Roewer, 1923 1690:Roewer, 1923 1685:Superfamily 1646:Phalangiidae 1628:Globipedidae 1617:Superfamily 1597:Superfamily 1542:Superfamily 1456: 1441:pre-Linnaean 1436: 1426: 1412: 1393:Rhynie chert 1383: 1361: 1355: 1348: 1322:Eugène Simon 1280: 1260: 1253:urban legend 1250: 1209: 1205: 1201: 1197: 1193: 1189: 1185: 1179: 1169: 1163: 1162: 1160:government. 1149: 1148: 1143: 1139: 1131: 1129: 1124: 1120: 1116: 1110: 1100: 1091:Bothriuridae 1083: 1079: 1071: 1067: 1064:Insidiatores 1063: 1059: 1053: 1044: 1031:stridulating 1028: 1025:Stridulation 1016: 1007: 1003: 999: 991: 969: 960: 943: 930:stridulation 915: 891: 887: 859:microhabitat 852: 845: 825: 816: 808: 792: 781: 760: 736: 708: 674: 660: 627: 625: 599:. A pair of 581:scent glands 578: 569: 544: 537:Gonyleptidae 530: 516: 513: 502: 479: 437: 414: 400: 367: 350:phylogenetic 347: 339: 332: 303: 299: 295: 291: 288:colloquially 275: 273: 210: 133: 29: 6692:iNaturalist 6550:Wikispecies 6124:(1): 1–46. 5142:J. Arachnol 5106:: 127–133. 5100:Anim. Behav 4294:"Opiliones" 4245:"Opiliones" 4191:Icaleptidae 4186:Guasiniidae 4168:Zalmoxoidea 4054:Gerdesiidae 3978:Epedanoidea 3963:Suthepiidae 3945:Assamioidea 3936:Grassatores 3887:Travuniidae 3734:Troguloidea 3719:Sabaconidae 3635:Neogoveidae 3397:Holothyrida 3361:Acariformes 3317:Acaromorpha 3288:Schizomida 3187:(scorpions) 3185:Scorpiones 3147:Palpigradi 3114:Chelicerata 3112:Subphylum: 2935:Sabaconidae 2858:Menge, 1854 2806:dentipalpis 2742:Haupt, 1956 2707:Lagerstätte 2558:Mazon Creek 2543:Mazon Creek 2473:, Scotland) 2334:basal clade 2287:The family 2254:Icaleptidae 2245:Guasiniidae 2218:Zalmoxoidea 2152:Simon, 1879 2143:Simon, 1879 2123:Simon, 1879 2057:Gerdesiidae 1947:Epedanoidea 1927:Suthepiidae 1900:Assamioidea 1891:Grassatores 1889:Infraorder 1849:Travuniidae 1825:Hadži, 1935 1807:Loman, 1900 1802:Infraorder 1766:Simon, 1872 1757:Simon, 1879 1745:Troguloidea 1725:Sabaconidae 1719:Simon, 1879 1710:Simon, 1879 1667:Simon, 1879 1658:Banks, 1893 1611:Banks, 1892 1602:Banks, 1892 1565:Shear, 1980 1556:Shear, 1980 1553:Neogoveidae 1547:Shear, 1980 1536:Shear, 1993 1522:Infraorder 1516:Shear, 1980 1502:Infraorder 1487:Simon, 1879 1473:Infraorder 1468:Simon, 1879 1447:Systematics 1103:troglobitic 1087:bonariensis 1068:Grassatores 1041:Retaliation 934:retaliation 928:, fleeing, 924:, bobbing, 880:predators. 863:leaf litter 832:aposematism 721:. Some are 662:Protolophus 555:sea spiders 541:Guasiniidae 523:Grassatores 463:opisthosoma 385:Description 191:Chelicerata 187:Subphylum: 41:410–0 6834:Harvestmen 6828:Categories 6177:Cladistics 6031:2012-12-21 5540:(1): 5–7. 5432:(1): 173. 4794:2015-05-29 4725:2007-08-27 4618:2023-07-16 4308:2024-07-10 4279:2017-11-29 4254:2024-07-10 4232:References 4201:Zalmoxidae 4039:Cosmetidae 4006:Tithaeidae 4001:Podoctidae 3991:Epedanidae 3953:Assamiidae 3933:Infraorder 3852:Infraorder 3844:Laniatores 3757:Trogulidae 3654:Infraorder 3624:Infraorder 3615:Pettalidae 3604:Infraorder 3579:Infraorder 3542:Arthropoda 3329:Ricinulei 3266:Amblypygi 3212:Solifugae 3177:Opiliones 3108:Arthropoda 3047:Quaternary 3034:Phalangium 3020:Pellobunus 2933:(Dyspnoi: 2890:(Dyspnoi: 2746:Trogulidae 2744:(Dyspnoi: 2365:ovipositor 2330:Laniatores 2272:Zalmoxidae 2087:Kury, 1994 2051:Kury, 2014 2033:Koch, 1839 2030:Cosmetidae 2015:Kury, 2003 1983:Tithaeidae 1974:Podoctidae 1956:Epedanidae 1909:Assamiidae 1894:Kury, 2002 1794:Laniatores 1781:Trogulidae 1544:Ogoveoidea 1513:Pettalidae 1271:chelicerae 1214:California 1085:Bothriurus 1072:Palpatores 951:Laniatores 940:Thanatosis 918:thanatosis 894:translucid 874:liverworts 849:camouflage 804:Gregarious 796:Laniatores 732:gregarines 723:scavengers 711:omnivorous 696:harvestman 681:tarsomeres 634:Trogulidae 613:copulation 593:book lungs 505:stomotheca 475:chelicerae 453:, and the 376:, and the 374:Tipuloidea 370:arthropods 335:Antarctica 324:Laniatores 296:harvesters 292:harvestmen 247:Laniatores 224:Suborders 181:Arthropoda 6570:Opiliones 6556:Opiliones 6526:Opiliones 6503:(fossils) 6475:Opiliones 6463:Opiliones 6388:April 16, 6315:1664-8021 6256:1314-2836 6199:1096-0031 6138:1096-3642 6102:1095-8312 5872:Research. 5562:0937-7409 5502:0161-8202 5446:0161-8202 5301:CiteSeerX 4992:2177-6199 4841:Behaviour 4638:Curr Biol 4550:0737-4038 4502:1063-5157 4196:Kimulidae 4130:Biantidae 4122:Samooidea 4089:Stygnidae 4044:Cranaidae 4029:Ampycidae 3782:Caddoidea 3724:Taracidae 3640:Ogoveidae 3590:Sironidae 3560:Opiliones 3548:Arachnida 3534:Kingdom: 3470:Haptopoda 3254:Pedipalpi 3243:(spiders) 3161:Dromopoda 3089:Arachnida 2986:Sironidae 2958:Oligocene 2948:(Eupnoi: 2912:Nemastoma 2899:Mitostoma 2873:(Eupnoi: 2865:Leiobunum 2832:(Eupnoi: 2811:(Eupnoi: 2782:Kimulidae 2750:Geiseltal 2711:Koonwarra 2530:, France) 2509:Commentry 2488:(Western 2399:Edinburgh 2381:E. sheari 2355:Paleozoic 2349:Paleozoic 2263:Kimulidae 2169:Biantidae 2160:Samooidea 2120:Stygnidae 2039:Cranaidae 2012:Ampycidae 1792:Suborder 1734:Taracidae 1675:Suborder 1599:Caddoidea 1587:Suborder 1562:Ogoveidae 1484:Sironidae 1463:Suborder 1423:Etymology 1409:Dromopoda 1405:solifuges 1397:scorpions 1386:arachnids 1374:Phylogeny 1267:crane fly 1237:Solifugae 1019:Leiobunum 977:Pholcidae 898:carapaces 854:Leiobunum 784:nocturnal 747:fossorial 737:Although 601:spiracles 509:pedipalps 455:posterior 290:known as 284:arachnids 276:Opiliones 262:Diversity 216:Sundevall 211:Opiliones 201:Arachnida 167:Kingdom: 161:Eukaryota 35:Opiliones 6601:BugGuide 6577:BioLib: 6535:Wikidata 6441:16024358 6333:25120562 6274:25425936 6215:85408627 6207:34772266 6146:84029705 6010:19495718 5932:Archived 5588:300–308. 5570:31558441 5510:83994309 5454:85822815 5331:10906916 5323:16273426 5200:26470886 5162:85621336 5120:54254258 5078:83850344 5035:85961182 4951:85846440 4943:40233818 4904:18019589 4861:49329266 4770:38401545 4712:85918640 4668:24726154 4568:35137183 4510:30917194 4461:21863011 4407:24726154 4135:Samoidae 3841:Suborder 3790:Caddidae 3770:Suborder 3676:Suborder 3656:(Indet.) 3568:Suborder 3540:Phylum: 3536:Animalia 3402:Ixodida 3296:Uropygi 3241:Araneae 3106:Phylum: 3102:Animalia 3100:Kingdom 3009:A. lawei 2928:claviger 2915:incertum 2868:longipes 2813:Caddidae 2792:silhavyi 2767:Samoidae 2738:longipes 2735:Trogulus 2721:Cenozoic 2715:Victoria 2696:Triassic 2595:Mesozoic 2490:Missouri 2452:Scotland 2385:H. argus 2369:tracheae 2345:Mesozoic 2341:Cenozoic 2319:Silurian 2178:Samoidae 1608:Caddidae 1390:Devonian 1277:Research 1265:and the 1257:venomous 1076:Ctenidae 1056:ozopores 1050:Chemical 983:Autotomy 957:Freezing 926:autotomy 922:freezing 640:Behavior 609:gonopore 597:tracheae 589:quinones 585:ozopores 498:antennae 494:medially 490:carapace 444:anterior 177:Phylum: 171:Animalia 157:Domain: 116:Holocene 112:Devonian 6811:1300999 6432:1599874 6324:4112917 6299:: 255. 6265:4238074 6064:, p.321 6018:9570512 5990:Bibcode 5542:Bibcode 5293:Bibcode 5208:1994257 5186:: 1–7. 5058:Bibcode 5015:Bibcode 4884:Bibcode 4646:Bibcode 4559:8845124 4439:Bibcode 4437:: 444. 4385:Bibcode 3679:Dyspnoi 3546:Class: 3460:Extinct 3404:(ticks) 3023:proavus 2956:, USA ( 2925:Sabacon 2826:ramiger 2754:Germany 2667:Myanmar 2648:Myanmar 2629:Myanmar 2610:Myanmar 2560:, U.S.) 2545:, U.S.) 2492:, U.S.) 2423:Permian 2270:Family 2261:Family 2252:Family 2243:Family 2234:Family 2225:Family 2205:Family 2185:Family 2176:Family 2167:Family 2147:Family 2127:Family 2118:Family 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Koch 1235:in the 1229:Chelate 1158:Spanish 1013:Fleeing 966:Bobbing 947:Dyspnoi 878:diurnal 867:epizoic 842:Crypsis 836:mimicry 828:crypsis 788:diurnal 777:instars 715:insects 673:A male 574:spiders 529:nests, 527:termite 525:) from 459:abdomen 451:prosoma 440:tagmata 359:Araneae 357:(order 355:spiders 320:Dyspnoi 242:Dyspnoi 207:Order: 197:Class: 6785:133884 6756:NZOR: 6645:1OPIOO 6541:Q19116 6439: 6429: 6358: 6331: 6321: 6313: 6272: 6262: 6254: 6213: 6205: 6197: 6144: 6136: 6100: 6016: 6008: 5911: 5637:87–99. 5568: 5560: 5508: 5500: 5452: 5444: 5373: 5329: 5321: 5303: 5229: 5206: 5198: 5160: 5118: 5076: 5033: 4990: 4949: 4941: 4902: 4859: 4818: 4768: 4710: 4666: 4591: 4566: 4556: 4548: 4508: 4500: 4459: 4405: 4349: 3773:Eupnoi 3557:Order 3128:Extant 3091:orders 2844:ovalis 2841:Opilio 2777:Kimula 2727:Eocene 2513:France 2448:Rhynie 2389:Eupnoi 1589:Eupnoi 1437:opilio 1403:, and 1233:chelae 1208:, and 1107:Brazil 972:Eupnoi 834:, and 800:Eupnoi 755:mating 559:Eupnoi 486:scutum 482:dorsal 442:, the 419:) body 348:Their 326:, and 316:Eupnoi 308:extant 237:Eupnoi 218:, 1833 110:Early 6806:WoRMS 6793:Plazi 6749:43271 6723:82753 6710:10452 6705:IRMNG 6697:47367 6658:15591 6593:24643 6580:16797 6211:S2CID 6142:S2CID 6025:(PDF) 6014:S2CID 5976:(PDF) 5566:S2CID 5506:S2CID 5450:S2CID 5327:S2CID 5204:S2CID 5158:S2CID 5116:S2CID 5074:S2CID 5031:S2CID 4947:S2CID 4939:JSTOR 4900:S2CID 4857:S2CID 4719:(PDF) 4708:S2CID 4686:(PDF) 4536:(2). 3343:Acari 3043:Italy 2803:Caddo 2769:?) — 2659:indet 2312:amber 1176:Texas 727:fecal 719:fungi 692:Mites 650:skink 605:tibia 471:venom 302:, or 280:order 6772:4428 6744:NCBI 6718:ITIS 6679:GBIF 6640:EPPO 6606:2405 6588:BOLD 6437:PMID 6390:2014 6356:ISBN 6329:PMID 6311:ISSN 6270:PMID 6252:ISSN 6203:PMID 6195:ISSN 6134:ISSN 6098:ISSN 6006:PMID 5909:ISBN 5558:ISSN 5498:ISSN 5442:ISSN 5371:ISBN 5319:PMID 5227:ISBN 5196:PMID 4988:ISSN 4816:ISBN 4766:PMID 4664:PMID 4589:ISBN 4564:PMID 4546:ISSN 4506:PMID 4498:ISSN 4457:PMID 4403:PMID 4347:ISBN 3280:s.l. 3039:Rome 2952:) — 2815:) — 2748:) — 2383:and 1168:and 1123:and 1101:All 949:and 872:and 652:tail 621:eggs 553:and 467:oval 274:The 49:PreꞒ 6731:NBN 6684:907 6632:249 6627:EoL 6619:3Q2 6614:CoL 6565:AFD 6427:PMC 6419:doi 6415:272 6319:PMC 6301:doi 6260:PMC 6242:doi 6185:doi 6126:doi 6122:174 6088:doi 6084:105 5998:doi 5550:doi 5490:doi 5434:doi 5311:doi 5260:doi 5188:doi 5184:121 5150:doi 5108:doi 5066:doi 5023:doi 4978:doi 4931:doi 4892:doi 4849:doi 4845:151 4758:doi 4700:doi 4654:doi 4554:PMC 4538:doi 4488:doi 4447:doi 4393:doi 3011:?) 2328:or 1251:An 1239:. 461:or 449:or 282:of 6830:: 6808:: 6795:: 6782:: 6769:: 6746:: 6733:: 6720:: 6707:: 6694:: 6681:: 6668:: 6655:: 6642:: 6629:: 6616:: 6603:: 6590:: 6567:: 6552:: 6537:: 6435:. 6425:. 6413:. 6407:. 6379:. 6350:. 6327:. 6317:. 6309:. 6295:. 6291:. 6268:. 6258:. 6250:. 6236:. 6232:. 6209:. 6201:. 6193:. 6181:31 6179:. 6175:. 6140:. 6132:. 6120:. 6096:. 6082:. 6078:. 6051:.) 6012:. 6004:. 5996:. 5986:96 5984:. 5978:. 5963:^ 5951:. 5826:^ 5814:^ 5809:4. 5783:^ 5771:^ 5746:^ 5702:^ 5672:^ 5642:^ 5629:^ 5578:^ 5564:. 5556:. 5548:. 5538:14 5536:. 5532:. 5518:^ 5504:. 5496:. 5486:40 5484:. 5480:. 5448:. 5440:. 5430:30 5428:. 5424:. 5394:^ 5339:^ 5325:. 5317:. 5309:. 5299:. 5289:31 5287:. 5274:^ 5256:16 5254:. 5250:. 5202:. 5194:. 5182:. 5170:^ 5156:. 5146:34 5144:. 5140:. 5128:^ 5114:. 5104:81 5102:. 5086:^ 5072:. 5064:. 5054:36 5052:. 5029:. 5021:. 5011:37 5009:. 4986:. 4974:13 4972:. 4968:. 4945:. 4937:. 4927:37 4925:. 4921:. 4898:. 4890:. 4880:13 4878:. 4855:. 4843:. 4764:. 4754:34 4752:. 4748:. 4734:^ 4706:. 4696:31 4694:. 4688:. 4662:. 4652:. 4642:24 4640:. 4636:. 4611:. 4562:. 4552:. 4544:. 4534:39 4532:. 4528:. 4504:. 4496:. 4484:68 4482:. 4478:. 4455:. 4445:. 4433:. 4429:. 4415:^ 4401:. 4391:. 4381:24 4379:. 4375:. 4361:^ 4341:. 4317:^ 4296:. 4272:. 4247:. 3041:, 2752:, 2729:. 2713:, 2669:, 2650:, 2631:, 2612:, 2511:, 2450:, 2351:. 2314:. 1419:. 1399:, 1370:. 1353:. 1314:, 1304:, 1204:, 1200:, 1196:, 1192:, 1188:, 1178:. 1127:. 1119:, 1115:, 932:, 920:, 908:. 838:. 830:, 802:. 753:, 734:. 623:. 337:. 322:, 318:, 314:, 298:, 294:, 286:, 114:– 99:Pg 43:Ma 6443:. 6421:: 6392:. 6364:. 6335:. 6303:: 6297:5 6276:. 6244:: 6238:2 6217:. 6187:: 6148:. 6128:: 6104:. 6090:: 6034:. 6000:: 5992:: 5957:. 5917:. 5572:. 5552:: 5544:: 5512:. 5492:: 5456:. 5436:: 5379:. 5333:. 5313:: 5295:: 5268:. 5262:: 5235:. 5210:. 5190:: 5164:. 5152:: 5122:. 5110:: 5080:. 5068:: 5060:: 5037:. 5025:: 5017:: 4994:. 4980:: 4953:. 4933:: 4906:. 4894:: 4886:: 4863:. 4851:: 4824:. 4797:. 4772:. 4760:: 4728:. 4702:: 4670:. 4656:: 4648:: 4621:. 4570:. 4540:: 4512:. 4490:: 4463:. 4449:: 4441:: 4435:2 4409:. 4395:: 4387:: 4355:. 4311:. 4282:. 4257:. 3518:e 3511:t 3504:v 3081:e 3074:t 3067:v 3049:) 3045:( 3007:( 2960:) 2515:) 2469:( 2454:) 2446:( 1089:( 632:( 583:( 535:( 521:( 405:) 251:† 104:N 94:K 89:J 84:T 79:P 74:C 69:D 64:S 59:O 54:Ꞓ 27:.

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