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
974:
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
774:
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
1045:
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
944:
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
5731:
DomĂnguez, 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
817:
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
761:
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
757:
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
729:
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
514:
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
380:
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
5890:
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
1457:
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
888:
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.
1004:
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.
1000:
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
846:
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
758:
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.
4523:
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
352:
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., MacĂas-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,
2309:
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
429:
428:
425:
5636:
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
344:
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.
427:
5282:
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?".
6115:
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)".
5972:
961:
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.
1009:
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.
5778:
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
5871:
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
5713:
Escalante, I., AlbĂn, 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.
5881:
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.
806:
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.
730:
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
6074:
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).
5762:
Mattoni, C.I., GarcĂa 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
1443:
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.
5178:
AlbĂn, A.; Toscano-Gadea, C. A. (2015). "Predation among armored arachnids: Bothriurus bonariensis (Scorpions, Bothriuridae) versus four species of harvestmen (Harvestmen, Gonyleptidae)".
5615:
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.
515:
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.
5658:
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.
438:
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
5606:
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.
2367:
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,
2317:
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
2964:
6158:
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.
4744:
Gainett, Guilherme; Klementz, Benjamin C.; Blaszczyk, Pola; Setton, Emily V.W.; Murayama, Gabriel P.; Willemart, Rodrigo; Gavish-Regev, Efrat; Sharma, Prashant P. (February 2024).
368:
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
2477:
5697:
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.
4783:
5833:
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.
5649:
Machado, G., Pomini, A.M., 2008. Chemical and behavioral defences of the neotropical harvestman Camarana flavipalpi (Arachnida: Opiliones). Biochem. Syst. Ecol. 36, 369–376.
2884:
6021:
5048:
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.).
770:
has evolved at least three times independently: once in the clade Progonyleptoidellinae + Caelopyginae, once in the Gonyleptinae, and once in the Heteropachylinae.
5799:
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.
603:
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 (200Ă— 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.
5466:
Machado, G., Vital, D.M., 2001. On the Occurrence of Epizoic Cyanobacteria and Liverworts on a Neotropical Harvestman (Arachnida: Opiliones) | BIOTROPICA
4682:
2978:
2805:
2737:
5722:
Huber, B.A., Eberhard, W.G., 1997. Courtship, copulation, and genital mechanics in Physocyclus globosus (Araneae, Pholcidae). Can. J. Zool. 75, 905–918.
5098:
da Silva Souza, E.; Willemart, R.H. (2011). "Harvest-ironman: heavy armature and not its defensive secretions, protects a harvestman against a spider".
2927:
1074:
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:
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.
6843:
3967:
1935:
1281:
Harvestmen are a scientifically neglected group. Description of new taxa has always been dependent on the activity of a few dedicated taxonomists.
1259:
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
480:
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,
5790:
Guffey, C., 1998. Leg Autotomy and Its Potential Fitness Costs for Two Species of Harvestmen (Arachnida, Opiliones). J. Arachnol. 26, 296–302.
5741:
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)".
4167:
4053:
3962:
2217:
2056:
1926:
790:
species are known, some of which have vivid patterns in yellow, green, and black with varied reddish and blackish mottling and reticulation.
5821:
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.
5597:
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
138:
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
6678:
4787:
2953:
2776:
1082:
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".
6717:
5764:
Primer caso de autotomĂa en escorpiones (Scorpiones: Buthidae). III Congreso Latinoamericano de AracnologĂa, Memorias y ResĂşmenes
2658:
1285:
described about a third (2,260) of today's known species from the 1910s to the 1950s, and published the landmark systematic work
2379:, which was though to form the sister group to Cyphophthalmi. However, recent reanalysis of harvestman phylogeny has shown that
1054:
Harvestmen are well known for being chemically protected. They exude strongly odored secretions from their scent glands, called
2709:
that has yielded many other terrestrial arachnids. An unnamed long-legged harvestman was reported from the Early Cretaceous of
5005:
Cáceres, N.C. (2002). "Food Habits and Seed Dispersal by the White-Eared Opossum, Didelphis albiventris, in Southern Brazil".
1001:
their legs, and leg loss reduces locomotion, speed, climbing ability, sensory perception, food detection, and territoriality.
6359:
5912:
5374:
4819:
4592:
4350:
1142:
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).
916:
Secondary defences allow for harvestmen to escape and survive from a predator after direct or indirect contact, including
5667:
Misslin, R., 2003. The defence system of fear: behaviour and neurocircuitry. Neurophysiol. Clin. Neurophysiol. 33, 55–66.
3516:
3079:
892:
Other harvestmen may exhibit mimicry to resemble other species' appearances. Some Gonyleptidae individuals that produce
6474:
6409:
1367:
876:
growing on their bodies that suggest potential benefits for camouflage against large backgrounds to avoid detection by
496:
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.
1269:, which are both also called a "daddy longlegs"). None of the known species of harvestmen have venom glands; their
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.
818:
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.
5861:
A Novel Class of Defensive Compounds in Harvestmen: Hydroxy-Îł-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).
4525:
2339:
Naturally, most finds are from comparatively recent times. More than 20 fossil species are known from the
970:
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
1062:
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, JesĂşs 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, TaĂs 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:Ă–zdikmen & Kury, 2007
2201:Ă–zdikmen & 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 & PĂ©rez, 2002
2230:Kury & PĂ©rez, 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:, JĂĽrgen Gruber,
1112:Giupponia chagasi
1097:Endangered status
902:MĂĽllerian 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:Ĺ ilhavĂ˝, 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: â€
2578:
2571:(Dyspnoi: â€
2563:
2548:
2533:
2518:
2505:Eotrogulidae
2503:(Dyspnoi: â€
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:HadĹľi, 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
2109:Family
2100:Family
2091:Family
2082:Family
2073:Family
2064:Family
2055:Family
2046:Family
2037:Family
2028:Family
2019:Family
2010:Family
2001:Family
1981:Family
1972:Family
1963:Family
1954:Family
1934:Family
1925:Family
1916:Family
1907:Family
1876:Family
1867:Family
1847:Family
1838:Family
1829:Family
1820:Family
1779:Family
1770:Family
1761:Family
1752:Family
1732:Family
1723:Family
1714:Family
1694:Family
1677:Dyspnoi
1662:Family
1653:Family
1644:Family
1635:Family
1626:Family
1606:Family
1574:Family
1560:Family
1551:Family
1531:Family
1511:Family
1491:Family
1482:Family
1312:L. 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:
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6016:
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5637:87–99.
5568:
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4508:
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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:Preęž’
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
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4654:doi
4554:PMC
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