668:
407:
999:
656:
160:
1253:
421:
137:
6481:
713:
576:
1236:
699:
6469:
681:
568:), 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
435:
1019:
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
985:
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
785:
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
1056:
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
955:
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
5742:
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
828:
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
772:
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
768:
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
740:
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
525:
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
391:
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
5901:
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
1468:
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
899:
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.
1015:
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.
1011:
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
857:
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
769:
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.
4534:
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
363:
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
1406:, 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
5598:
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,
2320:
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
440:
439:
436:
5647:
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.
441:
647:), 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.
5819:
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
355:
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.
438:
5293:
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?".
6126:
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)".
5983:
972:
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.
1020:
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.
5789:
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.
372:), 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
804:
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
5882:
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
5724:
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.
5892:
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.
476:. 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
5412:
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.
817:
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.
741:
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
6085:
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).
5773:
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
1454:
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.
5189:
AlbĂn, A.; Toscano-Gadea, C. A. (2015). "Predation among armored arachnids: Bothriurus bonariensis (Scorpions, Bothriuridae) versus four species of harvestmen (Harvestmen, Gonyleptidae)".
5626:
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.
526:
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
376:. 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.
5690:
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.
5669:
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.
449:
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
5617:
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.
2378:
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,
2328:
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
2975:
6169:
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.
4755:
Gainett, Guilherme; Klementz, Benjamin C.; Blaszczyk, Pola; Setton, Emily V.W.; Murayama, Gabriel P.; Willemart, Rodrigo; Gavish-Regev, Efrat; Sharma, Prashant P. (February 2024).
379:
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
2488:
5708:
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.
4794:
5844:
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.
5660:
Machado, G., Pomini, A.M., 2008. Chemical and behavioral defences of the neotropical harvestman Camarana flavipalpi (Arachnida: Opiliones). Biochem. Syst. Ecol. 36, 369–376.
2895:
6032:
5059:
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".
2952:
2770:
437:
4930:"Reproductive Behavior of Chavesincola inexpectabilis (Opiliones, Gonyleptidae) with Description of a New and Independently Evolved Case of Paternal Care in Harvestmen"
2802:
2612:
4348:
Glauco Machado, Ricardo Pinto-da-Rocha & Gonzalo Giribet (2007). "What are harvestmen?". In Ricardo Pinto-da-Rocha, Glauco Machado & Gonzalo Giribet (ed.).
781:
has evolved at least three times independently: once in the clade Progonyleptoidellinae + Caelopyginae, once in the Gonyleptinae, and once in the Heteropachylinae.
5810:
Macias-Ordonez, R., 1998. The mating system of Leiobunum vittatum Say 1821 (Arachnida: Opiliones: Palpatores): Resource defense polygyny in the striped harvestman.
2912:
2864:
2650:
2590:
2575:
412:
1426:, 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
2836:
2693:
2631:
2469:
1032:
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.
614:
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
528:
2560:
2545:
518:) 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
5853:
Eisner, T., Eisner, M., Siegler, M., 2005. Secret Weapons: Defenses of Insects, Spiders, Scorpions, and Other Many-legged Creatures. Harvard University Press.
2332:, 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.
1157:
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.
6087:"Evolutionary and biogeographical history of an ancient and global group of arachnids (Arachnida: Opiliones: Cyphophthalmi) with a new taxonomic arrangement"
2446:
1242:(pincer-like) chelicerae typical of harvestmen (200Ă— magnification); these chelicerae are homologous to chelicerae that take the form of fangs in spiders or
560:
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
6517:
6184:"Polyphyly of Caddoidea, reinstatement of the family Acropsopilionidae in Dyspnoi, and a revised classification system of Palpatores (Arachnida, Opiliones)"
4304:
3033:
3014:
3003:
2925:
2878:
2530:
3442:
5635:
Cokendolpher, J.C., 1987. Observations on the defensive behavior of a Neotropical Gonyleptidae (Arachnida, Opiliones). Revue Arachnologique, 7, pp.59–63.
5477:
Machado, G., Vital, D.M., 2001. On the Occurrence of Epizoic Cyanobacteria and Liverworts on a Neotropical Harvestman (Arachnida: Opiliones) | BIOTROPICA
4693:
2989:
2816:
2748:
5733:
Huber, B.A., Eberhard, W.G., 1997. Courtship, copulation, and genital mechanics in Physocyclus globosus (Araneae, Pholcidae). Can. J. Zool. 75, 905–918.
5109:
da Silva Souza, E.; Willemart, R.H. (2011). "Harvest-ironman: heavy armature and not its defensive secretions, protects a harvestman against a spider".
2938:
1085:
use substances like naphthoquinones, methyl- and ethyl-ketones. These secretions have successfully protected the harvestmen against wandering spiders (
503:. 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
4059:
2799:
2058:
5942:
5862:
Shultz, J.W., Pinto-da-Rocha, R., 2007. Morphology and functional anatomy. Harvest. Biol. Opiliones Harv. Univ. Press Camb. Mass. Lond. Engl. 14–61.
6854:
3978:
1946:
1292:
Harvestmen are a scientifically neglected group. Description of new taxa has always been dependent on the activity of a few dedicated taxonomists.
1270:
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
491:
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
6715:
4084:
3968:
3915:
3592:
2892:
2854:
2103:
1928:
1485:
4820:
990:) that are commonly mistaken for daddy long-legs (Opiliones) also exhibit this behavior when their webs are disturbed or even during courtship.
4044:
3996:
2481:
2031:
637:
Typical body length does not exceed 7 mm (0.28 in), and some species are smaller than 1 mm, although the largest known species,
5801:
Guffey, C., 1998. Leg Autotomy and Its Potential Fitness Costs for Two Species of Harvestmen (Arachnida, Opiliones). J. Arachnol. 26, 296–302.
5752:
Fleming, P.A., Muller, D., Bateman, P.W., 2007. Leave it all behind: a taxonomic perspective of autotomy in invertebrates. Biological Reviews.
1446:(1638–1712) by adopting Lister's term Opiliones for this order, known in Lister's days as "harvest spiders" or "shepherd spiders", from Latin
6754:
4885:
Machado, G.; Raimundo, R. L. G. (2001). "Parental investment and the evolution of subsocial behaviour in harvestmen (Arachnida: Opiliones)".
4178:
4064:
3973:
2228:
2067:
1937:
801:
species are known, some of which have vivid patterns in yellow, green, and black with varied reddish and blackish mottling and reticulation.
5832:
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.
5608:
Humphreys, R.K., Ruxton, G.D., 2018. A review of thanatosis (death feigning) as an anti-predator behaviour. Behav. Ecol. Sociobiol. 72, 22.
3030:
1554:
1302:(Harvestmen of the World) in 1923, with descriptions of all species known to that time. Other important taxonomists in this field include:
1028:
Individuals that are able to detect potential threats can flee rapidly from attack. This is seen with multiple long-legged species in the
149:
showing its body structure and long legs: one pair of eyes and broadly joined body tagma differentiate it from similar-looking arachnids.
6859:
5149:"The spider Enoploctenus cyclothorax (Araneae, Ctenidae) avoids preying on the harvestman Mischonyx cuspidatus (Opiliones, Gonyleptidae)"
4039:
2909:
2022:
911:. This may have an aposematic role by mimicking the colouration of glandular emissions of two other quinone-producing species. Mimicry (
1069:, that act as a shield against predators; this is the most effective defense they use which creates a strong and unpleasant taste. In
6387:
872:
used at each life stage. Many species have also been able to camouflage their bodies by covering with secretions and debris from the
6689:
4798:
2964:
2787:
1093:
ants. However, these chemical irritants are not able to prevent four species of harvestmen being preyed upon by the black scorpion
6058:. Ed. John Parker and Basil Hartley (1992). Colchester, Essex: Harley Books. pp. 26 & 30. (Translation of the Latin original,
2429:
was shown in 2004 to be a harvestman, after residing for almost one hundred years in its own arachnid order, the "Kustarachnida".
6728:
5775:
Primer caso de autotomĂa en escorpiones (Scorpiones: Buthidae). III Congreso Latinoamericano de AracnologĂa, Memorias y ResĂşmenes
2669:
1296:
described about a third (2,260) of today's known species from the 1910s to the 1950s, and published the landmark systematic work
2390:, which was though to form the sister group to Cyphophthalmi. However, recent reanalysis of harvestman phylogeny has shown that
1065:
Harvestmen are well known for being chemically protected. They exude strongly odored secretions from their scent glands, called
2720:
that has yielded many other terrestrial arachnids. An unnamed long-legged harvestman was reported from the Early Cretaceous of
5016:
Cáceres, N.C. (2002). "Food Habits and Seed Dispersal by the White-Eared Opossum, Didelphis albiventris, in Southern Brazil".
1012:
their legs, and leg loss reduces locomotion, speed, climbing ability, sensory perception, food detection, and territoriality.
6370:
5923:
5385:
4830:
4603:
4361:
1153:
is in rainforest patches in northwest Argentina which are in an area being dramatically destroyed by humans. The cave-living
1422:. The Opiliones have remained almost unchanged morphologically over a long period. Indeed, one species discovered in China,
6733:
6495:—A synoptic taxonomic arrangement of the order Opiliones, down to family-group level, including some photos of the families
1347:
2701:
dating from approximately 100 Mya. It has been suggested that this may be related to the Ortholasmatinae (Nemastomatidae).
927:
Secondary defences allow for harvestmen to escape and survive from a predator after direct or indirect contact, including
5678:
Misslin, R., 2003. The defence system of fear: behaviour and neurocircuitry. Neurophysiol. Clin. Neurophysiol. 33, 55–66.
3527:
3090:
903:
Other harvestmen may exhibit mimicry to resemble other species' appearances. Some Gonyleptidae individuals that produce
6485:
6420:
1378:
887:
growing on their bodies that suggest potential benefits for camouflage against large backgrounds to avoid detection by
507:
on the surface to form two plates beside each other. The second pair of legs is longer than the others and function as
4487:"A Critical Appraisal of the Placement of Xiphosura (Chelicerata) with Account of Known Sources of Phylogenetic Error"
2688:
etc. Bartel et al, 2023 report "These new records bring the total number of Burmese amber laniatorean species to ten"
5959:
5241:
5433:"A new species of pseudotrogulus roewer and assignment of the genus to the hernandariinae (opiliones, gonyleptidae)"
6816:
6668:
1284:
are not hollowed fangs but grasping claws that are typically very small and not strong enough to break human skin.
1280:, which are both also called a "daddy longlegs"). None of the known species of harvestmen have venom glands; their
1167:. It is included in the Catálogo Nacional de especies amenazadas (National catalog of threatened species) from the
6759:
2412:
in Scotland is almost 340 million years old. Its placement is rather uncertain, apart from it being a harvestman.
829:
chemical secretions. Some of these defences have been attributed and restricted to specific groups of harvestmen.
6681:
4312:
1120:. Four species of Opiliones are on the Brazilian national list of endangered species, all of them cave-dwelling:
317:. As of July 2024, over 6,650 species of harvestmen have been discovered worldwide, although the total number of
31:
6357:
Dunlop, Jason A. (2007). "Paleontology". In Ricardo Pinto-da-Rocha, Glauco Machado & Gonzalo Giribet (ed.).
5448:
4726:
1450:, "shepherd"; Lister characterized three species from England (although not formally describing them, being a
6741:
5489:"Epizoic cyanobacteria associated with a Neotropical harvestman (Opiliones: Sclerosomatidae) from Costa Rica"
904:
583:), showing the almost fused arrangement of abdomen and cephalothorax that distinguishes these arachnids from
5764:
Roth, V.D., Roth, B.M., 1984. review of appendotomy in spiders and other arachnids. Bull.-Br. Arachnol. Soc.
5872:
A Novel Class of Defensive Compounds in Harvestmen: Hydroxy-Îł-Lactones from the Phalangiid Egaenus convexus
4537:"Comprehensive Species Sampling and Sophisticated Algorithmic Approaches Refute the Monophyly of Arachnida"
1326:
504:
492:
465:
454:
6241:"World Checklist of Opiliones species (Arachnida). Part 1: Laniatores – Travunioidea and Triaenonychoidea"
5487:
Proud, Daniel N.; Wade, Ryan R.; Rock, Philip; Townsend, Victor R.; Chavez, Donald Jiménez (August 2012).
4536:
2350:
Naturally, most finds are from comparatively recent times. More than 20 fossil species are known from the
981:
To deflect attacks and enhance escape, long-legged species – commonly known as daddy long-legs – from the
6598:
6575:
5939:
5432:
4645:"A Paleozoic Stem Group to Mite Harvestmen Revealed through Integration of Phylogenetics and Development"
4384:"A Paleozoic Stem Group to Mite Harvestmen Revealed through Integration of Phylogenetics and Development"
1343:
6603:
6473:
2402:, after it was discovered that living daddy-longlegs have the same arrangement of eyes as the fossils.
17:
6821:
6808:
159:
6769:
6392:
4438:"Anatomically modern Carboniferous harvestmen demonstrate early cladogenesis and stasis in Opiliones"
598:) that secrete a peculiar-smelling fluid when disturbed. In some species, the fluid contains noxious
5316:
1306:
6849:
6492:
6362:
4353:
4280:
2304:
1367:
Early work on the developmental biology of Opiliones from the mid-20th century was resurrected by
4756:
3520:
3083:
35:
6795:
6504:
1104:). These secretions contain multiple volatile compounds that vary among individuals and clades.
814:
6528:
5311:
4701:
4619:
2922:
1360:
Since the 1990s, study of the biology and ecology of harvestmen has intensified, especially in
1073:
the scent glands release naphthoquinones, chloro-naphthoquinones and aliphatic methyl ketones,
1048:
and have also been shown to be used as a second line of defense when restrained by a predator.
623:
5913:
4694:"Third species of Guasiniidae (Opiliones, Laniatores) with comments on familial relationships"
4593:
6790:
1339:
1293:
1272:
1191:
774:
611:
6720:
5541:"Mimicry: imitative depiction of discharged defensive secretion on carapace of an opilionid"
4347:
6637:
6416:"First identifiable Mesozoic harvestman (Opiliones: Dyspnoi) from Cretaceous Burmese amber"
6000:
5552:
5303:
5068:
5025:
4894:
4656:
4486:
4449:
4395:
2725:
2379:
1463:
1439:
1390:
1045:
1003:
916:
667:
607:
226:
6782:
5699:
Field, L.H., Glasgow, S., 2001. The Biology of Wetas, King Crickets and Their Allies. CABI
4976:
4757:"Vestigial organs alter fossil placements in an ancient group of terrestrial chelicerates"
1227:
are being considered for listing as endangered species, but as yet receive no protection.
912:
8:
6844:
6566:
5991:
5421:
Edgar, A.L., 1971. Studies on the biology and ecology of Michigan Phalangida (Opiliones).
4795:"SENCKENBERG world of biodiversity | About us | Communications | Newsroom"
2496:
2477:
1141:
Several Opiliones in Argentina appear to be vulnerable, if not endangered. These include
837:
Primary defences help the harvestmen avoid encountering a potential predator and include
820:
Harvestmen clean their legs after eating by drawing each leg in turn through their jaws.
753:
6004:
5871:
5556:
5307:
5072:
5029:
4898:
4660:
4453:
4399:
6442:
6415:
6334:
6299:
6275:
6240:
6221:
6152:
6024:
5576:
5516:
5460:
5337:
5214:
5168:
5126:
5084:
5041:
4957:
4949:
4910:
4867:
4718:
4569:
3716:
3698:
3513:
3076:
2813:
2538:
1717:
1697:
1368:
426:
406:
154:
145:
1185:
are listed as endangered species in the United States. Both are from caves in central
1077:
use nitrogen-containing substances, terpenes, aliphatic ketones, and phenolics, while
868:
are multiple species with cryptic colouration that changes over ontogeny to match the
6777:
6624:
6447:
6366:
6339:
6321:
6280:
6262:
6213:
6205:
6144:
6108:
6103:
6086:
6016:
5919:
5568:
5508:
5452:
5381:
5329:
5237:
5206:
4998:
4826:
4776:
4674:
4599:
4574:
4556:
4516:
4508:
4467:
4413:
4382:
Garwood, Russell J.; Sharma, Prashant P.; Dunlop, Jason A.; Giribet, Gonzalo (2014).
4357:
3925:
3762:
3752:
3724:
3706:
3402:
2507:
2299:
1879:
1782:
1764:
1726:
1706:
1451:
1373:
1122:
932:
686:
556:
542:
508:
401:
6225:
6156:
5580:
5520:
5464:
5341:
5172:
5130:
5088:
5045:
4961:
4914:
4871:
4850:(Arachnida: Opiliones): from premating struggles to solicitous tactile engagement".
4722:
964:
suborders, with individuals becoming rigid with legs either retracted or stretched.
915:) occurring between Brazilian harvestmen that resemble others could be explained by
6437:
6429:
6329:
6311:
6270:
6252:
6195:
6136:
6098:
6028:
6008:
5560:
5500:
5444:
5321:
5270:
5218:
5198:
5160:
5118:
5076:
5033:
4988:
4977:"Reproductive Behavior in Harvestman (Arachnida): Mating Systems and Parental Care"
4941:
4902:
4859:
4768:
4710:
4664:
4598:, vol. 5: Harvester ant to leaf-cutting ant, Marshall Cavendish, p. 263,
4564:
4548:
4498:
4457:
4403:
3907:
3485:
2710:
1870:
1316:
1312:
1175:
1086:
639:
599:
5981:
5148:
5122:
4906:
6629:
6580:
5946:
5202:
4436:
Garwood, Russell J.; Dunlop, Jason A.; Giribet, Gonzalo; Sutton, Mark D. (2011).
4191:
3930:
3887:
3838:
3655:
3617:
2885:
2781:
2713:
2598:
2583:
2374:
is known from two specimens, one a female, the other a male. The female bears an
2309:
2246:
1888:
1841:
1674:
1543:
1514:
1427:
1353:
998:
884:
749:
627:
450:
290:
6746:
1332:
6676:
6663:
5540:
4150:
4114:
4089:
3892:
3882:
3757:
3637:
3490:
3394:
3384:
3379:
3239:
3214:
3149:
2982:
2902:
2454:
2197:
2150:
2112:
1850:
1832:
1773:
1534:
1411:
1181:
1164:
1146:
928:
798:
655:
561:
6511:
6012:
5564:
5325:
5080:
5037:
4993:
4863:
4846:
Fowler-Finn, K.D.; Triana, E.; Miller, O.G. (2014). "Mating in the harvestman
4772:
4669:
4644:
4408:
4383:
2717:
2347:, and the latter having probably diverged from the Dyspnoi more than 300 Mya.
344:
Representatives of each extant suborder can be found on all continents except
6838:
6325:
6316:
6266:
6209:
6148:
6112:
5572:
5512:
5456:
5002:
4560:
4552:
4512:
4160:
4122:
4034:
4026:
3833:
3823:
3810:
3673:
3605:
3581:
2960:
2833:
2698:
2681:
2662:
2643:
2624:
2416:
2387:
2336:
2208:
2159:
2013:
2004:
1665:
1647:
1629:
1586:
1503:
1475:
1443:
1361:
1252:
1243:
1239:
1057:
spiders by causing injury, allowing the harvestman to escape from predation.
880:
782:
778:
761:
631:
457:
373:
360:
338:
322:
263:
242:
84:
6498:
6170:
5275:
5258:
4503:
4485:
Ballesteros, JesĂşs A; Sharma, Prashant P (2019-11-01). Halanych, Ken (ed.).
3505:
2343:
much older than 50 million years are known, despite the former presenting a
6551:
6451:
6433:
6343:
6284:
6217:
6020:
5964:
5333:
5210:
4780:
4678:
4578:
4520:
4471:
4417:
4186:
4168:
4104:
4094:
4079:
4074:
4069:
4006:
3920:
3874:
3865:
3828:
3818:
3420:
2844:
2827:
2515:
2458:
2237:
2217:
2139:
2121:
2094:
2085:
2076:
1975:
1823:
1814:
1656:
1638:
1403:
1263:
1101:
1041:
940:
547:
352:
318:
272:
1149:, the highest peak in the Sierras Chicas chain (provincia de Cordoba) and
420:
6702:
6590:
6560:
4533:
4201:
4196:
3988:
3955:
3946:
3897:
3744:
3729:
3645:
3407:
3371:
3124:
2945:
2568:
2553:
2344:
2264:
2255:
1957:
1910:
1901:
1859:
1755:
1735:
1563:
1116:
species (of all animal taxa) are considered to be at least threatened in
1113:
944:
873:
842:
672:
591:
554:. However, recent work studying the embryonic development of the species
551:
533:
481:
473:
298:
201:
59:
6257:
4953:
4929:
1469:
the placement and even monophyly of several taxa are still in question.
321:
species may exceed 10,000. The order Opiliones includes five suborders:
6239:
Kury, Adriano B.; Mendes, Amanda Cruz; Souza, Daniele R. (2014-11-05).
4462:
4437:
4211:
4049:
4016:
4011:
4001:
3963:
3854:
3767:
3625:
3480:
3298:
3157:
3057:
3044:
2756:
2375:
2340:
2282:
2040:
1993:
1984:
1966:
1919:
1804:
1791:
1523:
1281:
1277:
1224:
1095:
986:
individuals, which are all trembling at the same time. Cellar spiders (
961:
888:
859:
806:
721:
706:
644:
603:
565:
515:
485:
384:
345:
334:
257:
136:
104:
69:
6616:
6200:
6183:
6140:
5488:
4975:
Machado, Glauco; Requena, Gustavo S.; Buzatto, Bruno A. (2009-12-06).
4945:
2423:
species are currently grouped as Dyspnoi, they look more like Eupnoi.
6707:
6650:
5984:"Harvestmen (Arachnida: Opiliones) from the Middle Jurassic of China"
5504:
5164:
4206:
4140:
4132:
4099:
4054:
3792:
3734:
3650:
3600:
3558:
3552:
3339:
3276:
3222:
3170:
3118:
2996:
2968:
2875:
2792:
2760:
2721:
2519:
2409:
2359:
2312:
is sometimes misplaced in the Phalangioidea. It is not a harvestman.
2273:
2179:
2170:
2130:
2049:
1744:
1609:
1572:
1494:
1419:
1377:
as a model system for the study of arachnid comparative genomics and
1247:
1029:
987:
864:
794:
757:
742:
733:
712:
691:
480:
structure. Other differences include the fact that Opiliones have no
388:
380:
191:
171:
109:
53:
6522:
6501:—Diagnostic photographs and information on North American harvestmen
550:) from caves, most species of Cyphophthalmi, and all species of the
6694:
6655:
6611:
6545:
6480:
4714:
4145:
3800:
3264:
3195:
3099:
3068:
2823:
2777:
2745:
2706:
2500:
2462:
2355:
2351:
2329:
2188:
1618:
1415:
1407:
1400:
1396:
1235:
936:
908:
877:
619:
575:
519:
500:
351:
Well-preserved fossils have been found in the 400-million-year-old
294:
211:
126:
122:
99:
94:
79:
74:
64:
30:
For other arthropods called "daddy longlegs", and other uses, see
3689:
3306:
2935:
2764:
2677:
2658:
2639:
2620:
2433:
1687:
1322:
1066:
957:
869:
846:
838:
760:
species in the suborder Cyphophthalmi, where the males deposit a
595:
537:
469:
461:
369:
330:
252:
114:
89:
5539:
González, Andrés; Rossini, Carmen; Eisner, Thomas (2004-03-01).
5259:"Determining the fitness consequences of antipredation behavior"
4620:"4-Eyed Daddy Longlegs Helps Explain Arachnid Evolution (Video)"
6468:
3783:
3546:
3251:
3112:
2851:
2737:
2523:
2399:
1599:
1117:
982:
810:
790:
to reach maturity, with most known species having six instars.
787:
765:
725:
698:
584:
569:
365:
326:
247:
181:
6507:—Diagnostic photographs and information on European harvestmen
4754:
4260:
Kury, A. et al. (2023). WCO-Lite: World Catalogue of Opiliones
876:
found in their environments. Some hard-bodied harvestmen have
511:
or feelers. In short-legged species, this may not be obvious.
6803:
5380:. Cambridge, MA: Harvard University Press. pp. 374–399.
5292:
3352:
3053:
2382:, and no median eyes. Together with the 305-million-year-old
2322:
1267:
1223:
from around springs and other restricted habitats of central
1186:
1168:
737:
729:
660:
615:
496:
6642:
5915:
The Earwig's Tail: a modern bestiary of multi-legged legends
5234:
Defence in Animals : a Survey of Anti-predator Defences
4309:
Joel Hallan's Biology Catalog (Archive version. August 2017)
680:
6300:"A revised dated phylogeny of the arachnid order Opiliones"
6084:
5982:
Diying Huang, Paul A. Selden & Jason A. Dunlop (2009).
5449:
10.1636/0161-8202(2002)030[0173:ansopr]2.0.co;2
5430:
3412:
3049:
2436:
are possibly harvestmen, but these are not well preserved.
1040:
Multiple species within the Laniatores and Dyspnoi possess
702:
477:
5146:
4642:
4435:
4381:
1258:
Opilionid chelae shown in context to the rest of the body.
6388:"Ancient four-eyed wonder resolves daddy longleg mystery"
956:
a potential predator and is commonly observed within the
5376:
Gnaspini, P.; Hara, M. R. (2007). "Defense mechanisms".
4643:
Garwood, RJ; Sharma, PP; Dunlop, JA; Giribet, G (2014).
6518:
Joel Hallan's Biology Catalog (Archive only before 2017
5431:
Leandro Firmo, Carlos; Pinto-da-Rocha, Ricardo (2002).
5108:
4845:
3010:(Eupnoi: Sclerosomatidae) — Florissant, USA (Oligocene)
694:(the many small segments making up the end of each leg)
392:
better observe their wandering flocks from a distance.
4255:
2697:, a long-legged Dyspnoi with large eyes, was found in
5538:
5486:
5058:
4974:
2419:
of North America and Europe. While the two described
3064:
736:, feeding upon dead organisms, bird dung, and other
2705:Currently, no fossil harvestmen are known from the
1356:, Adriano Brilhante Kury, Tone Novak (21st century)
797:and colored in hues of brown, although a number of
445:
Harvestmen (Opiliones sp.) filmed in Hesse, Germany
5743:in the genus Prionostemma. J. Arachnol. 44, 76–82.
5188:
4692:Pinto-da-Rocha, Ricardo; Kury, Adriano B. (2003).
4691:
6125:
4484:
2736:Unless otherwise noted, all species are from the
6836:
6491:Adriano Kury: National Museum of Rio de Janeiro
2277:Pérez González, Kury & Alonso-Zarazaga, 2007
6238:
5902:western North America | Journal of Arachnology.
5147:Willemart, R.H.; Pellegatti-Franco, F. (2006).
4927:
4884:
4591:
2415:From about 300 Mya, several finds are from the
1081:use alkylated phenolics and benzoquinones, and
590:Harvestmen have a pair of prosomatic defensive
499:, which in most such species is fused with the
6298:Sharma, Prashant P.; Giribet, Gonzalo (2014).
4617:
3567:
862:when they remain motionless. Within the genus
6413:
6297:
6182:Groh, Selina; Giribet, Gonzalo (2015-06-01).
5905:
5256:
3535:
3521:
3084:
5375:
2586:) — Upper Carboniferous (Commentary, France)
1297:
1251:
6414:Giribet, Gonzalo; Dunlop, Jason A. (2005).
6181:
4928:Nazareth, TaĂs M.; Machado, Glauco (2009).
2601:) — Upper Carboniferous (Mazon Creek, U.S.)
4585:
3528:
3514:
3091:
3077:
894:
630:, unlike other arachnids. All species lay
135:
6441:
6333:
6315:
6274:
6256:
6199:
6171:https://dx.doi.org/10.1098/rspb.2016.2340
6129:Zoological Journal of the Linnean Society
6102:
6091:Biological Journal of the Linnean Society
5918:. Harvard University Press. p. 143.
5911:
5315:
5274:
4992:
4668:
4568:
4502:
4461:
4407:
1438:The Swedish naturalist and arachnologist
907:secretions have orange markings on their
5977:
5975:
3040:(Laniatores: Samoidae) — Dominican amber
2965:Florissant Fossil Beds National Monument
2931:(Dyspnoi: Nemastomatidae) — Baltic amber
2918:(Dyspnoi: Nemastomatidae) — Baltic amber
2809:(Laniatores: Samoidae) — Dominican amber
2709:. So far, they are also absent from the
1338:James C. Cokendolpher, Raymond Forster,
1234:
997:
823:
711:
697:
679:
666:
654:
574:
433:
419:
405:
6855:Extant Early Devonian first appearances
6073:Curious Facts in the History of Insects
5957:
5912:Berenbaum, May R. (30 September 2009).
5231:
5015:
4377:
4375:
4373:
4343:
4341:
4339:
4337:
4335:
4333:
4331:
4329:
4248:
2537:(Tetrophthalmi) — Upper Carboniferous (
1266:claims that the harvestman is the most
1163:has only been found in one cave in the
14:
6837:
6385:
6356:
5840:
5838:
5828:
5826:
5797:
5795:
5785:
5783:
5760:
5758:
5720:
5718:
5716:
5714:
5686:
5684:
5371:
4818:
4750:
4748:
4746:
4431:
4429:
4427:
3048:species (Eupnoi: Phalangiidae) — near
2871:(Eupnoi: Phalangiidae?) — Baltic amber
2860:(Eupnoi: Phalangiidae?) — Baltic amber
622:on the ventral cephalothorax, and the
6527:
6526:
6056:Martin Lister's English Spiders, 1678
5972:
5656:
5654:
5643:
5641:
5594:
5592:
5590:
5534:
5532:
5530:
5408:
5406:
5369:
5367:
5365:
5363:
5361:
5359:
5357:
5355:
5353:
5351:
5288:
5286:
5257:Lind, Johan; Cresswell, Will (2005).
5184:
5182:
5142:
5140:
5104:
5102:
5100:
5098:
4297:
3509:
3469:
3137:
3072:
922:
383:, the crane flies of the superfamily
34:. For the novel by Jean Webster, see
6809:022087FF-3A6B-980B-4FDC-84C4FD85F944
6770:be640fe4-ffd8-4b4d-a2bf-9323c3b71ff0
6682:535a87e2-e40b-4209-9156-f7f119df3b16
6359:Harvestmen: the Biology of Opiliones
5378:Harvestmen: The Biology of Opiliones
4618:Blaszczak-Boxe, Agata (2014-04-10).
4370:
4350:Harvestmen: the Biology of Opiliones
4326:
3098:
2728:, Australia, which may be a Eupnoi.
2439:
2398:are in fact members of the suborder
1590:Karaman, Mitov & Snegovaya, 2024
1107:
728:and all kinds of plant material and
5835:
5823:
5792:
5780:
5755:
5711:
5681:
4743:
4535:(2022-02-03). Teeling, Emma (ed.).
4424:
1442:(1801–1875) honored the naturalist
1418:; these four orders form the clade
832:
777:, and cleaning the eggs regularly.
24:
6860:Taxa named by Carl Jakob Sundevall
6421:Proceedings of the Royal Society B
5651:
5638:
5587:
5527:
5403:
5348:
5283:
5179:
5137:
5095:
1379:evolutionary-developmental biology
809:, and more than 70,000 in certain
752:species do occur, most harvestmen
25:
6871:
6461:
6386:Tepper, Fabien (April 11, 2014).
3065:Further internal navigation links
1195:from a cave in central Texas and
6479:
6467:
6104:10.1111/j.1095-8312.2011.01774.x
4887:Ethology Ecology & Evolution
4595:Insects and spiders of the world
4278:
3450:are possibly paraphyletic groups
2567:(Eupnoi) — Upper Carboniferous (
2552:(Eupnoi) — Upper Carboniferous (
2315:
1335:, William Sørensen (1880s–1890s)
1230:
1145:, which is found only on top of
1089:), wolf spiders (Lycosidae) and
716:Gregarious behavior in Opiliones
690:, showing the long legs and the
279:5 suborders, > 6,650 species
158:
57:
6407:
6379:
6350:
6291:
6232:
6175:
6163:
6119:
6078:
6065:
6049:
5951:
5932:
5895:
5886:
5876:
5865:
5856:
5847:
5813:
5804:
5767:
5746:
5736:
5727:
5702:
5693:
5672:
5663:
5629:
5620:
5611:
5602:
5480:
5471:
5424:
5415:
5394:
5250:
5225:
5061:Stud. Neotropical Fauna Environ
5052:
5018:Stud. Neotropical Fauna Environ
5009:
4968:
4921:
4878:
4839:
4812:
4787:
4685:
4636:
4611:
4541:Molecular Biology and Evolution
2976:Proholoscotolemon nemastomoides
2807:Cokendolpher & Poinar, 1998
1845:Derkarabetian & Hedin, 2018
1348:Cândido Firmino de Mello-Leitão
1035:
488:, so pose no danger to humans.
32:Daddy longlegs (disambiguation)
5958:Perkins, Sid (June 23, 2009).
4825:. Princeton University Press.
4819:Cowles, Jillian (2018-06-12).
4527:
4478:
4272:
2358:, and at least seven from the
1863:Absolon & Kratochvil, 1932
1827:Absolon & Kratochvil, 1932
1457:
1143:Pachyloidellus fulvigranulatus
1051:
395:
13:
1:
5960:"Long-lasting daddy longlegs"
5123:10.1016/j.anbehav.2010.09.023
4907:10.1080/08927014.2001.9522780
4592:Joyce Tavolacci, ed. (2003),
4281:"Classification of Opiliones"
4242:
3026:— Florissant, USA (Oligocene)
2308:) from underground waters in
1642:Kury & Cokendolpher, 2020
950:
289:(formerly Phalangida) are an
5203:10.1016/j.beproc.2015.10.003
3441:Classification is based on
2489:Echinopustulus samuelnelsoni
2365:
1433:
1384:
1340:Clarence and Marie Goodnight
1327:Tord Tamerlan Teodor Thorell
522:and the first pair of legs.
341:, which were named in 2014.
7:
6512:The Rhynie Chert Harvestmen
6505:Harvestman: Order Opiliones
6499:Harvestman: Order Opiliones
6493:Classification of Opiliones
5296:Journal of Chemical Ecology
3470:
3301:(shorttailed whipscorpions)
2731:
2605:
1691:Hansen & Sørensen, 1904
1603:Hansen & Sørensen, 1904
1507:Hansen & Sørensen, 1904
1344:Reginald Frederick Lawrence
1287:
1060:
1046:intraspecific communication
993:
967:
650:
628:male Opiliones have a penis
424:North European harvestman (
10:
6876:
4934:The Journal of Arachnology
3138:
2841:(Koch & Berendt, 1854)
1461:
1388:
1044:organs, which are used as
1023:
976:
852:
399:
29:
6535:
6393:Christian Science Monitor
6245:Biodiversity Data Journal
6013:10.1007/s00114-009-0556-3
5565:10.1007/s00049-003-0252-2
5326:10.1007/s10886-005-7611-0
5081:10.1076/snfe.36.2.85.2138
5038:10.1076/snfe.37.2.97.8582
4994:10.4257/oeco.2009.1301.05
4864:10.1163/1568539x-00003209
4773:10.1016/j.cub.2024.02.011
4670:10.1016/j.cub.2014.03.039
4409:10.1016/j.cub.2014.03.039
4285:www.museunacional.ufrj.br
4177:
4159:
4131:
4113:
4025:
3987:
3954:
3943:
3906:
3873:
3862:
3851:
3809:
3791:
3780:
3743:
3715:
3697:
3686:
3664:
3634:
3614:
3589:
3578:
3541:
3536:Extant Opiliones Families
3476:
3465:
3437:
3393:
3370:
3351:
3335:
3326:
3288:
3279:(tailless whip scorpions)
3272:
3263:
3247:
3238:
3207:
3180:
3169:
3148:
3144:
3133:
3107:
2953:Petrunkevitchiana oculata
2771:Philacarus hispaniolensis
2518:) — Upper Carboniferous (
2370:The 410-million-year-old
2221:Ă–zdikmen & Kury, 2007
2212:Ă–zdikmen & Kury, 2007
2035:Kury & Carvalho, 2020
1299:Die Weberknechte der Erde
1007:with several missing legs
947:and chemical secretions.
724:, eating primarily small
618:of the legs. They have a
278:
271:
239:
234:
155:Scientific classification
153:
143:
134:
45:
6510:University of Aberdeen:
6363:Harvard University Press
6317:10.3389/fgene.2014.00255
5400:Cokendolpher pers. comm.
4354:Harvard University Press
2676:(Insidiatores indet.) —
2613:Burmalomanius circularis
2386:, it forms the suborder
2305:Stygophalangium karamani
1997:Sharma and Giribet, 2011
1979:Sharma and Giribet, 2011
1932:Sharma and Giribet, 2011
1132:Pachylospeleus strinatii
5938:The Spider Myths Site:
2408:from East Kirkton near
1395:Harvestmen are ancient
1201:Microcina edgewoodensis
895:Aposematism and mimicry
572:(true daddy-longlegs).
514:The feeding apparatus (
36:Daddy-Long-Legs (novel)
6434:10.1098/rspb.2005.3063
5493:Journal of Arachnology
5437:Journal of Arachnology
4702:Journal of Arachnology
4553:10.1093/molbev/msac021
2865:Cheiromachus coriaceus
2651:Mesodibunus tourinhoae
2591:Nemastomoides longipes
2576:Nemastomoides elaveris
2432:Some fossils from the
2268:Kury & PĂ©rez, 2002
2241:Kury & PĂ©rez, 2003
1810:(about 4,200 species)
1605:(about 1,800 species)
1307:Pierre André Latreille
1298:
1259:
1256:
1136:Spaeleoleptes spaeleus
1008:
786:four to eight nymphal
717:
709:
695:
677:
664:
606:, and breathe through
587:
446:
431:
417:
413:Pachyloidellus goliath
6791:Paleobiology Database
6304:Frontiers in Genetics
5276:10.1093/beheco/ari075
5236:. New York: Longman.
4504:10.1093/sysbio/syz011
4442:Nature Communications
3160:(microwhip scorpions)
3024:(Petrunkevitch, 1922)
2957:(Petrunkevitch, 1922)
2791:species (Laniatores:
2694:Halitherses grimaldii
2632:Petroburma tarsomeria
2550:(Petrunkevitch, 1913)
2470:Brigantibunum listoni
2406:Brigantibunum listoni
2259:Gonzalez-Sponga, 1997
1294:Carl Friedrich Roewer
1273:Pholcus phalangioides
1255:
1238:
1192:Texella cokendolpheri
1155:Picunchenops spelaeus
1001:
824:Antipredator defences
715:
701:
683:
676:sp. cleaning its legs
670:
658:
579:A harvestman (a male
578:
529:Caecobunus termitarum
444:
423:
409:
6677:Fauna Europaea (new)
6476:at Wikimedia Commons
6365:. pp. 247–265.
6060:Tractatus de Araneis
5232:Edmunds, M. (1974).
4858:(12–13): 1663–1686.
3342:(hooded tickspiders)
2999:) — Bitterfeld amber
2561:Kustarachne tenuipes
2546:Kustarachne longipes
2427:Kustarachne tenuipes
1693:(about 400 species)
1582:Infraorder (indet).
1481:(about 200 species)
1464:Harvestman phylogeny
1440:Carl Jakob Sundevall
1391:Harvestman phylogeny
1151:Pachyloides borellii
1004:Rilaena triangularis
917:convergent evolution
756:. Except from small
659:Harvestman eating a
410:Chilean harvestman (
6428:(1567): 1007–1013.
6258:10.3897/BDJ.2.e4094
6005:2009NW.....96..955H
5992:Naturwissenschaften
5557:2004Checo..14....5G
5308:2005JCEco..31.2519M
5073:2001SNFE...36...85C
5030:2002SNFE...37...97C
4981:Oecologia Australis
4899:2001EtEcE..13..133M
4767:(6): 1258–1270.e5.
4661:2014CBio...24.1017G
4454:2011NatCo...2..444G
4400:2014CBio...24.1017G
2795:) — Dominican amber
2674:Bartel et al, 2023
2655:Bartel et al, 2023
2636:Bartel et al, 2023
2617:Bartel et al, 2023
2595:Petrunkevitch, 1913
2497:Upper Carboniferous
2478:Early Carboniferous
2447:Eophalangium sheari
2372:Eophalangium sheari
1399:. Fossils from the
1091:Formica exsectoides
602:. They do not have
5945:2007-07-14 at the
5263:Behavioral Ecology
4848:Lieobunum vittatum
4491:Systematic Biology
4463:10.1038/ncomms1458
3717:Ischyropsalidoidea
3699:Acropsopilionoidea
3215:Pseudoscorpionida
3038:Cokendolpher, 1987
3015:Amauropilio lacoei
3004:Amauropilio atavus
2539:Montceau-les-Mines
2531:Hastocularis argus
2453:(Tetrophthalmi) —
2384:Hastocularis argus
2335:No fossils of the
1718:Ischyropsalidoidea
1698:Acropsopilionoidea
1424:Mesobunus martensi
1371:, who established
1369:Prashant P. Sharma
1346:, Jochen Martens,
1260:
1257:
1221:Microcina tiburona
1009:
923:Secondary defences
775:egg-eating females
754:reproduce sexually
718:
710:
696:
678:
665:
588:
495:shield called the
447:
432:
427:Leiobunum rotundum
418:
146:Hadrobunus grandis
27:Order of arachnids
6832:
6831:
6778:Open Tree of Life
6529:Taxon identifiers
6472:Media related to
6372:978-0-674-02343-7
6201:10.1111/cla.12087
6141:10.1111/zoj.12225
5925:978-0-674-05356-4
5387:978-0-674-02343-7
5302:(11): 2519–2539.
4946:10.1636/ST08-32.1
4832:978-1-4008-9018-7
4822:Amazing Arachnids
4605:978-0-7614-7334-3
4363:978-0-674-02343-7
4356:. pp. 1–13.
4279:Kury, Adriano B.
4236:
4235:
4232:
4231:
4228:
4227:
4224:
4223:
4220:
4219:
3939:
3938:
3926:Synthetonychiidae
3847:
3846:
3776:
3775:
3763:Nipponopsalididae
3753:Dicranolasmatidae
3725:Ischyropsalididae
3707:Acropsopilionidae
3682:
3681:
3503:
3502:
3499:
3498:
3461:
3460:
3457:
3456:
3433:
3432:
3429:
3428:
3416:
3403:Opilioacariformes
3363:
3343:
3322:
3321:
3318:
3317:
3310:
3302:
3280:
3255:
3234:
3233:
3226:
3218:
3217:(pseudoscorpions)
3199:
3191:
3161:
3039:
3025:
3009:
3008:(Cockerell, 1907)
2994:
2990:Siro platypedibus
2980:
2958:
2943:
2930:
2917:
2900:
2883:
2870:
2859:
2842:
2821:
2808:
2775:
2754:
2675:
2656:
2638:(Petrobunidae) —
2637:
2618:
2596:
2581:
2566:
2551:
2536:
2513:
2508:Eotrogulus fayoli
2494:
2475:
2452:
2440:Described species
2354:, three from the
2300:Stygophalangiidae
2287:
2278:
2269:
2260:
2251:
2242:
2233:
2222:
2213:
2202:
2193:
2184:
2175:
2164:
2155:
2144:
2135:
2126:
2117:
2108:
2099:
2090:
2081:
2072:
2063:
2054:
2045:
2036:
2027:
2018:
2009:
1998:
1989:
1980:
1971:
1962:
1951:
1942:
1933:
1924:
1915:
1906:
1893:
1884:
1880:Synthetonychiidae
1875:
1864:
1855:
1846:
1837:
1828:
1819:
1809:
1796:
1787:
1783:Nipponopsalididae
1778:
1769:
1765:Dicranolasmatidae
1760:
1749:
1740:
1731:
1727:Ischyropsalididae
1722:
1711:
1707:Acropsopilionidae
1702:
1692:
1679:
1670:
1661:
1652:
1643:
1634:
1623:
1614:
1604:
1591:
1577:
1568:
1559:
1548:
1539:
1528:
1519:
1508:
1499:
1490:
1480:
1374:Phalangium opilio
1342:, JĂĽrgen Gruber,
1123:Giupponia chagasi
1108:Endangered status
913:MĂĽllerian mimicry
793:Most species are
720:Many species are
687:Phalangium opilio
581:Phalangium opilio
557:Phalangium opilio
543:Giupponia chagasi
442:
402:Opiliones anatomy
283:
282:
230:
129:
16:(Redirected from
6867:
6825:
6824:
6812:
6811:
6799:
6798:
6786:
6785:
6773:
6772:
6763:
6762:
6750:
6749:
6747:NBNSYS0000160574
6737:
6736:
6724:
6723:
6711:
6710:
6698:
6697:
6685:
6684:
6672:
6671:
6659:
6658:
6646:
6645:
6633:
6632:
6620:
6619:
6607:
6606:
6594:
6593:
6584:
6583:
6571:
6570:
6569:
6556:
6555:
6554:
6524:
6523:
6484:Data related to
6483:
6471:
6456:
6455:
6445:
6411:
6405:
6404:
6402:
6400:
6383:
6377:
6376:
6354:
6348:
6347:
6337:
6319:
6295:
6289:
6288:
6278:
6260:
6236:
6230:
6229:
6203:
6179:
6173:
6167:
6161:
6160:
6123:
6117:
6116:
6106:
6082:
6076:
6069:
6063:
6053:
6047:
6046:
6044:
6043:
6037:
6031:. Archived from
5988:
5979:
5970:
5969:
5955:
5949:
5940:"Daddy-Longlegs"
5936:
5930:
5929:
5909:
5903:
5899:
5893:
5890:
5884:
5880:
5874:
5869:
5863:
5860:
5854:
5851:
5845:
5842:
5833:
5830:
5821:
5817:
5811:
5808:
5802:
5799:
5790:
5787:
5778:
5771:
5765:
5762:
5753:
5750:
5744:
5740:
5734:
5731:
5725:
5722:
5709:
5706:
5700:
5697:
5691:
5688:
5679:
5676:
5670:
5667:
5661:
5658:
5649:
5645:
5636:
5633:
5627:
5624:
5618:
5615:
5609:
5606:
5600:
5596:
5585:
5584:
5536:
5525:
5524:
5505:10.1636/b11-24.1
5484:
5478:
5475:
5469:
5468:
5428:
5422:
5419:
5413:
5410:
5401:
5398:
5392:
5391:
5373:
5346:
5345:
5319:
5290:
5281:
5280:
5278:
5254:
5248:
5247:
5229:
5223:
5222:
5191:Behav. Processes
5186:
5177:
5176:
5165:10.1636/S05-70.1
5144:
5135:
5134:
5106:
5093:
5092:
5056:
5050:
5049:
5013:
5007:
5006:
4996:
4972:
4966:
4965:
4925:
4919:
4918:
4882:
4876:
4875:
4843:
4837:
4836:
4816:
4810:
4809:
4807:
4806:
4797:. Archived from
4791:
4785:
4784:
4752:
4741:
4740:
4738:
4737:
4731:
4725:. Archived from
4698:
4689:
4683:
4682:
4672:
4640:
4634:
4633:
4631:
4630:
4615:
4609:
4608:
4589:
4583:
4582:
4572:
4531:
4525:
4524:
4506:
4482:
4476:
4475:
4465:
4433:
4422:
4421:
4411:
4394:(9): 1017–1023.
4379:
4368:
4367:
4345:
4324:
4323:
4321:
4320:
4311:. Archived from
4301:
4295:
4294:
4292:
4291:
4276:
4270:
4269:
4267:
4266:
4252:
3952:
3951:
3908:Triaenonychoidea
3871:
3870:
3860:
3859:
3789:
3788:
3695:
3694:
3587:
3586:
3576:
3575:
3565:
3564:
3530:
3523:
3516:
3507:
3506:
3486:Phalangiotarbida
3467:
3466:
3449:
3414:
3368:
3367:
3362:
3361:
3357:
3355:
3341:
3333:
3332:
3329:
3308:
3300:
3278:
3270:
3269:
3253:
3245:
3244:
3224:
3216:
3197:
3189:
3178:
3177:
3173:
3159:
3146:
3145:
3135:
3134:
3093:
3086:
3079:
3070:
3069:
3037:
3023:
3007:
2995:(Cyphophthalmi:
2993:
2985:) — Baltic amber
2979:
2956:
2948:) — Baltic amber
2942:
2929:
2916:
2905:) — Baltic amber
2899:
2888:) — Baltic amber
2882:
2868:
2858:
2847:) — Baltic amber
2840:
2820:
2806:
2774:
2752:
2711:Lower Cretaceous
2673:
2654:
2635:
2616:
2594:
2579:
2564:
2549:
2534:
2511:
2492:
2473:
2450:
2285:
2276:
2267:
2258:
2249:
2240:
2231:
2220:
2211:
2200:
2191:
2182:
2173:
2162:
2153:
2142:
2133:
2124:
2115:
2106:
2097:
2088:
2079:
2071:Bragagnolo, 2015
2070:
2061:
2052:
2043:
2034:
2025:
2016:
2007:
1996:
1987:
1978:
1969:
1960:
1949:
1940:
1931:
1922:
1913:
1904:
1891:
1882:
1873:
1871:Triaenonychoidea
1862:
1853:
1844:
1835:
1826:
1817:
1807:
1794:
1785:
1776:
1767:
1758:
1748:Schönhofer, 2013
1747:
1738:
1729:
1720:
1709:
1700:
1690:
1677:
1668:
1659:
1650:
1641:
1632:
1621:
1612:
1602:
1589:
1575:
1566:
1557:
1546:
1537:
1526:
1517:
1506:
1497:
1488:
1478:
1317:Maximilian Perty
1313:Carl Ludwig Koch
1301:
1176:Texella reddelli
1161:Maiorerus randoi
833:Primary defences
640:Trogulus torosus
484:glands in their
443:
225:
163:
162:
139:
120:
119:
56:
49:Temporal range:
43:
42:
21:
6875:
6874:
6870:
6869:
6868:
6866:
6865:
6864:
6850:Arachnid orders
6835:
6834:
6833:
6828:
6820:
6815:
6807:
6802:
6794:
6789:
6781:
6776:
6768:
6766:
6758:
6753:
6745:
6740:
6732:
6727:
6719:
6714:
6706:
6701:
6693:
6688:
6680:
6675:
6667:
6662:
6654:
6649:
6641:
6636:
6628:
6623:
6615:
6610:
6602:
6597:
6589:
6587:
6579:
6574:
6565:
6564:
6559:
6550:
6549:
6544:
6531:
6464:
6459:
6412:
6408:
6398:
6396:
6384:
6380:
6373:
6355:
6351:
6296:
6292:
6237:
6233:
6180:
6176:
6168:
6164:
6124:
6120:
6083:
6079:
6070:
6066:
6054:
6050:
6041:
6039:
6035:
5986:
5980:
5973:
5956:
5952:
5947:Wayback Machine
5937:
5933:
5926:
5910:
5906:
5900:
5896:
5891:
5887:
5881:
5877:
5870:
5866:
5861:
5857:
5852:
5848:
5843:
5836:
5831:
5824:
5818:
5814:
5809:
5805:
5800:
5793:
5788:
5781:
5772:
5768:
5763:
5756:
5751:
5747:
5741:
5737:
5732:
5728:
5723:
5712:
5707:
5703:
5698:
5694:
5689:
5682:
5677:
5673:
5668:
5664:
5659:
5652:
5646:
5639:
5634:
5630:
5625:
5621:
5616:
5612:
5607:
5603:
5597:
5588:
5537:
5528:
5485:
5481:
5476:
5472:
5429:
5425:
5420:
5416:
5411:
5404:
5399:
5395:
5388:
5374:
5349:
5317:10.1.1.384.1362
5291:
5284:
5255:
5251:
5244:
5230:
5226:
5187:
5180:
5145:
5138:
5107:
5096:
5057:
5053:
5014:
5010:
4973:
4969:
4926:
4922:
4883:
4879:
4844:
4840:
4833:
4817:
4813:
4804:
4802:
4793:
4792:
4788:
4761:Current Biology
4753:
4744:
4735:
4733:
4729:
4696:
4690:
4686:
4641:
4637:
4628:
4626:
4624:livescience.com
4616:
4612:
4606:
4590:
4586:
4532:
4528:
4483:
4479:
4434:
4425:
4388:Current Biology
4380:
4371:
4364:
4346:
4327:
4318:
4316:
4303:
4302:
4298:
4289:
4287:
4277:
4273:
4264:
4262:
4254:
4253:
4249:
4245:
4239:
4237:
4216:
4192:Fissiphalliidae
4173:
4155:
4127:
4109:
4060:Cryptogeobiidae
4021:
3983:
3945:
3935:
3931:Triaenonychidae
3902:
3888:Cryptomastridae
3864:
3853:
3843:
3839:Sclerosomatidae
3805:
3782:
3772:
3739:
3711:
3688:
3678:
3666:
3660:
3656:Troglosironidae
3636:
3630:
3618:Scopulophthalmi
3616:
3610:
3591:
3580:
3569:
3563:
3537:
3534:
3504:
3495:
3472:
3453:
3447:
3425:
3389:
3359:
3358:
3356:
3353:
3347:
3327:
3314:
3284:
3259:
3230:
3225:(camel spiders)
3203:
3171:
3165:
3140:
3129:
3103:
3097:
3067:
2886:Sclerosomatidae
2800:Hummelinckiolus
2782:Dominican amber
2734:
2714:Crato Formation
2657:(Epedanidae) —
2619:(Podoctidae) —
2608:
2599:Nemastomoididae
2584:Nemastomoididae
2442:
2368:
2318:
2310:North Macedonia
2296:
2247:Fissiphalliidae
2080:Sundevall, 1833
2059:Cryptogeobiidae
2008:Sundevall, 1833
1889:Triaenonychidae
1842:Cryptomastridae
1795:Sundevall, 1833
1759:Sundevall, 1833
1675:Sclerosomatidae
1660:Latreille, 1802
1633:Latreille, 1802
1544:Troglosironidae
1515:Scopulophthalmi
1466:
1460:
1436:
1428:Sclerosomatidae
1412:pseudoscorpions
1393:
1387:
1354:Gonzalo Giribet
1290:
1233:
1209:Microcina jungi
1110:
1063:
1054:
1038:
1026:
996:
979:
970:
953:
925:
897:
855:
835:
826:
750:parthenogenetic
705:parasitising a
653:
562:horseshoe crabs
434:
404:
398:
311:harvest spiders
224:
157:
130:
118:
117:
112:
107:
102:
97:
92:
87:
82:
77:
72:
67:
62:
51:
50:
47:
39:
28:
23:
22:
15:
12:
11:
5:
6873:
6863:
6862:
6857:
6852:
6847:
6830:
6829:
6827:
6826:
6813:
6800:
6787:
6774:
6764:
6751:
6738:
6725:
6712:
6699:
6686:
6673:
6664:Fauna Europaea
6660:
6647:
6634:
6621:
6608:
6595:
6585:
6572:
6557:
6541:
6539:
6533:
6532:
6521:
6520:
6515:
6508:
6502:
6496:
6489:
6488:at Wikispecies
6477:
6463:
6462:External links
6460:
6458:
6457:
6406:
6378:
6371:
6349:
6290:
6231:
6194:(3): 277–290.
6174:
6162:
6118:
6077:
6064:
6048:
5999:(8): 955–962.
5971:
5950:
5931:
5924:
5904:
5894:
5885:
5875:
5864:
5855:
5846:
5834:
5822:
5812:
5803:
5791:
5779:
5766:
5754:
5745:
5735:
5726:
5710:
5701:
5692:
5680:
5671:
5662:
5650:
5637:
5628:
5619:
5610:
5601:
5586:
5526:
5499:(2): 259–261.
5479:
5470:
5423:
5414:
5402:
5393:
5386:
5347:
5282:
5269:(5): 945–956.
5249:
5242:
5224:
5178:
5159:(3): 649–652.
5136:
5094:
5051:
5008:
4967:
4940:(2): 127–134.
4920:
4893:(2): 133–150.
4877:
4838:
4831:
4811:
4786:
4742:
4715:10.1636/H02-59
4709:(3): 394–399.
4684:
4655:(9): 1017–23.
4635:
4610:
4604:
4584:
4526:
4497:(6): 896–917.
4477:
4423:
4369:
4362:
4325:
4296:
4271:
4246:
4244:
4241:
4234:
4233:
4230:
4229:
4226:
4225:
4222:
4221:
4218:
4217:
4215:
4214:
4209:
4204:
4199:
4194:
4189:
4183:
4181:
4175:
4174:
4172:
4171:
4165:
4163:
4157:
4156:
4154:
4153:
4151:Stygnommatidae
4148:
4143:
4137:
4135:
4129:
4128:
4126:
4125:
4119:
4117:
4115:Phalangodoidea
4111:
4110:
4108:
4107:
4102:
4097:
4092:
4090:Otilioleptidae
4087:
4082:
4077:
4072:
4067:
4062:
4057:
4052:
4047:
4042:
4037:
4031:
4029:
4023:
4022:
4020:
4019:
4014:
4009:
4004:
3999:
3993:
3991:
3985:
3984:
3982:
3981:
3979:Trionyxellidae
3976:
3971:
3966:
3960:
3958:
3949:
3941:
3940:
3937:
3936:
3934:
3933:
3928:
3923:
3918:
3912:
3910:
3904:
3903:
3901:
3900:
3895:
3893:Paranonychidae
3890:
3885:
3883:Cladonychiidae
3879:
3877:
3868:
3857:
3849:
3848:
3845:
3844:
3842:
3841:
3836:
3831:
3826:
3821:
3815:
3813:
3807:
3806:
3804:
3803:
3797:
3795:
3786:
3778:
3777:
3774:
3773:
3771:
3770:
3765:
3760:
3758:Nemastomatidae
3755:
3749:
3747:
3741:
3740:
3738:
3737:
3732:
3727:
3721:
3719:
3713:
3712:
3710:
3709:
3703:
3701:
3692:
3684:
3683:
3680:
3679:
3677:
3676:
3670:
3668:
3662:
3661:
3659:
3658:
3653:
3648:
3642:
3640:
3638:Sternophthalmi
3632:
3631:
3629:
3628:
3622:
3620:
3612:
3611:
3609:
3608:
3603:
3597:
3595:
3584:
3573:
3562:
3561:
3555:
3549:
3542:
3539:
3538:
3533:
3532:
3525:
3518:
3510:
3501:
3500:
3497:
3496:
3494:
3493:
3491:Trigonotarbida
3488:
3483:
3477:
3474:
3473:
3463:
3462:
3459:
3458:
3455:
3454:
3452:
3451:
3448:Items in green
3445:
3438:
3435:
3434:
3431:
3430:
3427:
3426:
3424:
3423:
3418:
3410:
3405:
3399:
3397:
3395:Parasitiformes
3391:
3390:
3388:
3387:
3385:Trombidiformes
3382:
3380:Sarcoptiformes
3376:
3374:
3365:
3360:(ticks, mites)
3349:
3348:
3346:
3345:
3336:
3330:
3324:
3323:
3320:
3319:
3316:
3315:
3313:
3312:
3304:
3295:
3293:
3286:
3285:
3283:
3282:
3273:
3267:
3261:
3260:
3258:
3257:
3248:
3242:
3240:Tetrapulmonata
3236:
3235:
3232:
3231:
3229:
3228:
3220:
3211:
3209:
3205:
3204:
3202:
3201:
3193:
3184:
3182:
3175:
3167:
3166:
3164:
3163:
3154:
3152:
3150:Incertae sedis
3142:
3141:
3131:
3130:
3128:
3127:
3121:
3115:
3108:
3105:
3104:
3096:
3095:
3088:
3081:
3073:
3066:
3063:
3062:
3061:
3041:
3027:
3011:
3000:
2986:
2983:Cladonychiidae
2972:
2949:
2932:
2919:
2906:
2903:Nemastomatidae
2889:
2872:
2861:
2848:
2830:
2810:
2796:
2784:
2767:
2733:
2730:
2703:
2702:
2686:
2685:
2666:
2647:
2628:
2607:
2604:
2603:
2602:
2587:
2580:Thevenin, 1901
2572:
2557:
2542:
2527:
2512:Thevenin, 1901
2504:
2485:
2466:
2455:Early Devonian
2441:
2438:
2367:
2364:
2317:
2314:
2302:(one species,
2295:
2294:
2293:
2292:
2291:
2290:
2289:
2288:
2286:Sørensen, 1886
2279:
2270:
2261:
2252:
2243:
2232:Sørensen, 1886
2225:
2224:
2223:
2205:
2204:
2203:
2198:Stygnommatidae
2194:
2192:Sørensen, 1886
2185:
2174:Sørensen, 1886
2167:
2166:
2165:
2151:Phalangodoidea
2147:
2146:
2145:
2143:Sørensen, 1932
2136:
2127:
2118:
2113:Otilioleptidae
2109:
2100:
2091:
2082:
2073:
2064:
2055:
2046:
2037:
2028:
2019:
2001:
2000:
1999:
1990:
1981:
1972:
1970:Sørensen, 1886
1961:Sørensen, 1886
1954:
1953:
1952:
1947:Trionyxellidae
1943:
1934:
1925:
1923:Sørensen, 1884
1914:Sørensen, 1884
1898:
1897:
1896:
1895:
1894:
1892:Sørensen, 1886
1885:
1874:Sørensen, 1886
1867:
1866:
1865:
1856:
1851:Paranonychidae
1847:
1838:
1833:Cladonychiidae
1801:
1800:
1799:
1798:
1797:
1788:
1779:
1774:Nemastomatidae
1770:
1752:
1751:
1750:
1741:
1732:
1714:
1713:
1712:
1684:
1683:
1682:
1681:
1680:
1671:
1662:
1653:
1651:Lawrence, 1931
1644:
1626:
1625:
1624:
1596:
1595:
1594:
1593:
1592:
1580:
1579:
1578:
1569:
1551:
1550:
1549:
1535:Sternophthalmi
1531:
1530:
1529:
1511:
1510:
1509:
1500:
1471:
1462:Main article:
1459:
1456:
1435:
1432:
1389:Main article:
1386:
1383:
1358:
1357:
1351:
1350:(20th century)
1336:
1330:
1320:
1310:
1309:(18th century)
1289:
1286:
1232:
1229:
1217:Microcina lumi
1213:Microcina leei
1205:Microcina homi
1197:Calicina minor
1182:Texella reyesi
1165:Canary Islands
1147:Cerro Uritorco
1128:Iandumoema uai
1109:
1106:
1062:
1059:
1053:
1050:
1037:
1034:
1025:
1022:
995:
992:
978:
975:
969:
966:
952:
949:
924:
921:
896:
893:
854:
851:
834:
831:
825:
822:
652:
649:
400:Main article:
397:
394:
389:cellar spiders
315:daddy longlegs
281:
280:
276:
275:
269:
268:
267:
266:
260:
255:
250:
245:
237:
236:
232:
231:
219:
215:
214:
209:
205:
204:
199:
195:
194:
189:
185:
184:
179:
175:
174:
169:
165:
164:
151:
150:
141:
140:
132:
131:
113:
108:
103:
98:
93:
88:
83:
78:
73:
68:
63:
58:
48:
26:
9:
6:
4:
3:
2:
6872:
6861:
6858:
6856:
6853:
6851:
6848:
6846:
6843:
6842:
6840:
6823:
6818:
6814:
6810:
6805:
6801:
6797:
6792:
6788:
6784:
6779:
6775:
6771:
6765:
6761:
6756:
6752:
6748:
6743:
6739:
6735:
6730:
6726:
6722:
6717:
6713:
6709:
6704:
6700:
6696:
6691:
6687:
6683:
6678:
6674:
6670:
6665:
6661:
6657:
6652:
6648:
6644:
6639:
6635:
6631:
6626:
6622:
6618:
6613:
6609:
6605:
6600:
6596:
6592:
6586:
6582:
6577:
6573:
6568:
6562:
6558:
6553:
6547:
6543:
6542:
6540:
6538:
6534:
6530:
6525:
6519:
6516:
6513:
6509:
6506:
6503:
6500:
6497:
6494:
6490:
6487:
6482:
6478:
6475:
6470:
6466:
6465:
6453:
6449:
6444:
6439:
6435:
6431:
6427:
6423:
6422:
6417:
6410:
6395:
6394:
6389:
6382:
6374:
6368:
6364:
6360:
6353:
6345:
6341:
6336:
6331:
6327:
6323:
6318:
6313:
6309:
6305:
6301:
6294:
6286:
6282:
6277:
6272:
6268:
6264:
6259:
6254:
6250:
6246:
6242:
6235:
6227:
6223:
6219:
6215:
6211:
6207:
6202:
6197:
6193:
6189:
6185:
6178:
6172:
6166:
6158:
6154:
6150:
6146:
6142:
6138:
6134:
6130:
6122:
6114:
6110:
6105:
6100:
6097:(1): 92–130.
6096:
6092:
6088:
6081:
6074:
6071:Frank Cowan,
6068:
6061:
6057:
6052:
6038:on 2016-03-05
6034:
6030:
6026:
6022:
6018:
6014:
6010:
6006:
6002:
5998:
5994:
5993:
5985:
5978:
5976:
5967:
5966:
5961:
5954:
5948:
5944:
5941:
5935:
5927:
5921:
5917:
5916:
5908:
5898:
5889:
5879:
5873:
5868:
5859:
5850:
5841:
5839:
5829:
5827:
5816:
5807:
5798:
5796:
5786:
5784:
5776:
5770:
5761:
5759:
5749:
5739:
5730:
5721:
5719:
5717:
5715:
5705:
5696:
5687:
5685:
5675:
5666:
5657:
5655:
5644:
5642:
5632:
5623:
5614:
5605:
5595:
5593:
5591:
5582:
5578:
5574:
5570:
5566:
5562:
5558:
5554:
5550:
5546:
5542:
5535:
5533:
5531:
5522:
5518:
5514:
5510:
5506:
5502:
5498:
5494:
5490:
5483:
5474:
5466:
5462:
5458:
5454:
5450:
5446:
5442:
5438:
5434:
5427:
5418:
5409:
5407:
5397:
5389:
5383:
5379:
5372:
5370:
5368:
5366:
5364:
5362:
5360:
5358:
5356:
5354:
5352:
5343:
5339:
5335:
5331:
5327:
5323:
5318:
5313:
5309:
5305:
5301:
5297:
5289:
5287:
5277:
5272:
5268:
5264:
5260:
5253:
5245:
5243:0-582-44132-3
5239:
5235:
5228:
5220:
5216:
5212:
5208:
5204:
5200:
5196:
5192:
5185:
5183:
5174:
5170:
5166:
5162:
5158:
5154:
5150:
5143:
5141:
5132:
5128:
5124:
5120:
5116:
5112:
5105:
5103:
5101:
5099:
5090:
5086:
5082:
5078:
5074:
5070:
5066:
5062:
5055:
5047:
5043:
5039:
5035:
5031:
5027:
5024:(2): 97–104.
5023:
5019:
5012:
5004:
5000:
4995:
4990:
4986:
4982:
4978:
4971:
4963:
4959:
4955:
4951:
4947:
4943:
4939:
4935:
4931:
4924:
4916:
4912:
4908:
4904:
4900:
4896:
4892:
4888:
4881:
4873:
4869:
4865:
4861:
4857:
4853:
4849:
4842:
4834:
4828:
4824:
4823:
4815:
4801:on 2015-11-19
4800:
4796:
4790:
4782:
4778:
4774:
4770:
4766:
4762:
4758:
4751:
4749:
4747:
4732:on 2016-03-28
4728:
4724:
4720:
4716:
4712:
4708:
4704:
4703:
4695:
4688:
4680:
4676:
4671:
4666:
4662:
4658:
4654:
4650:
4646:
4639:
4625:
4621:
4614:
4607:
4601:
4597:
4596:
4588:
4580:
4576:
4571:
4566:
4562:
4558:
4554:
4550:
4546:
4542:
4538:
4530:
4522:
4518:
4514:
4510:
4505:
4500:
4496:
4492:
4488:
4481:
4473:
4469:
4464:
4459:
4455:
4451:
4447:
4443:
4439:
4432:
4430:
4428:
4419:
4415:
4410:
4405:
4401:
4397:
4393:
4389:
4385:
4378:
4376:
4374:
4365:
4359:
4355:
4351:
4344:
4342:
4340:
4338:
4336:
4334:
4332:
4330:
4315:on 2017-08-04
4314:
4310:
4306:
4300:
4286:
4282:
4275:
4261:
4257:
4251:
4247:
4240:
4213:
4210:
4208:
4205:
4203:
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4190:
4188:
4185:
4184:
4182:
4180:
4176:
4170:
4167:
4166:
4164:
4162:
4161:Sandokanoidea
4158:
4152:
4149:
4147:
4144:
4142:
4139:
4138:
4136:
4134:
4130:
4124:
4123:Phalangodidae
4121:
4120:
4118:
4116:
4112:
4106:
4103:
4101:
4098:
4096:
4093:
4091:
4088:
4086:
4085:Nomoclastidae
4083:
4081:
4078:
4076:
4073:
4071:
4068:
4066:
4063:
4061:
4058:
4056:
4053:
4051:
4048:
4046:
4043:
4041:
4038:
4036:
4035:Agoristenidae
4033:
4032:
4030:
4028:
4027:Gonyleptoidea
4024:
4018:
4015:
4013:
4010:
4008:
4005:
4003:
4000:
3998:
3995:
3994:
3992:
3990:
3986:
3980:
3977:
3975:
3972:
3970:
3969:Pyramidopidae
3967:
3965:
3962:
3961:
3959:
3957:
3953:
3950:
3948:
3942:
3932:
3929:
3927:
3924:
3922:
3919:
3917:
3916:Buemarinoidae
3914:
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3911:
3909:
3905:
3899:
3896:
3894:
3891:
3889:
3886:
3884:
3881:
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3878:
3876:
3872:
3869:
3867:
3861:
3858:
3856:
3850:
3840:
3837:
3835:
3834:Protolophidae
3832:
3830:
3827:
3825:
3824:Neopilionidae
3822:
3820:
3817:
3816:
3814:
3812:
3811:Phalangioidea
3808:
3802:
3799:
3798:
3796:
3794:
3790:
3787:
3785:
3779:
3769:
3766:
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3714:
3708:
3705:
3704:
3702:
3700:
3696:
3693:
3691:
3685:
3675:
3674:Parasironidae
3672:
3671:
3669:
3663:
3657:
3654:
3652:
3649:
3647:
3644:
3643:
3641:
3639:
3633:
3627:
3624:
3623:
3621:
3619:
3613:
3607:
3606:Stylocellidae
3604:
3602:
3599:
3598:
3596:
3594:
3593:Boreophthalmi
3588:
3585:
3583:
3582:Cyphophthalmi
3577:
3574:
3572:
3566:
3560:
3556:
3554:
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3492:
3489:
3487:
3484:
3482:
3479:
3478:
3475:
3468:
3464:
3446:
3444:
3443:Shultz (2007)
3440:
3439:
3436:
3422:
3419:
3417:
3411:
3409:
3406:
3404:
3401:
3400:
3398:
3396:
3392:
3386:
3383:
3381:
3378:
3377:
3375:
3373:
3369:
3366:
3364:
3350:
3344:
3338:
3337:
3334:
3331:
3325:
3311:
3309:(vinegaroons)
3305:
3303:
3297:
3296:
3294:
3292:
3289:Thelyphonida
3287:
3281:
3275:
3274:
3271:
3268:
3266:
3262:
3256:
3250:
3249:
3246:
3243:
3241:
3237:
3227:
3221:
3219:
3213:
3212:
3210:
3206:
3200:
3194:
3192:
3186:
3185:
3183:
3179:
3176:
3174:
3168:
3162:
3156:
3155:
3153:
3151:
3147:
3143:
3136:
3132:
3126:
3122:
3120:
3116:
3114:
3110:
3109:
3106:
3101:
3094:
3089:
3087:
3082:
3080:
3075:
3074:
3071:
3059:
3055:
3051:
3047:
3046:
3042:
3036:
3035:
3032:
3028:
3021:
3017:
3016:
3012:
3006:
3005:
3001:
2998:
2992:
2991:
2987:
2984:
2981:(Laniatores:
2978:
2977:
2973:
2970:
2966:
2962:
2961:Phalangioidea
2955:
2954:
2950:
2947:
2941:
2940:
2937:
2933:
2928:
2927:
2924:
2920:
2915:
2914:
2911:
2907:
2904:
2898:
2897:
2894:
2893:Histricostoma
2890:
2887:
2881:
2880:
2877:
2873:
2867:
2866:
2862:
2857:
2856:
2853:
2849:
2846:
2839:
2838:
2835:
2834:Dicranopalpus
2831:
2829:
2825:
2819:
2818:
2815:
2811:
2805:
2804:
2801:
2797:
2794:
2790:
2789:
2785:
2783:
2779:
2776:(Laniatores:
2773:
2772:
2768:
2766:
2762:
2758:
2751:
2750:
2747:
2743:
2742:
2741:
2739:
2729:
2727:
2723:
2719:
2716:of Brazil, a
2715:
2712:
2708:
2700:
2699:Burmese amber
2696:
2695:
2691:
2690:
2689:
2683:
2682:Burmese amber
2679:
2672:
2671:
2667:
2664:
2663:Burmese amber
2660:
2653:
2652:
2648:
2645:
2644:Burmese amber
2641:
2634:
2633:
2629:
2626:
2625:Burmese amber
2622:
2615:
2614:
2610:
2609:
2600:
2593:
2592:
2588:
2585:
2578:
2577:
2573:
2570:
2565:Scudder, 1890
2563:
2562:
2558:
2555:
2548:
2547:
2543:
2540:
2535:Garwood, 2014
2533:
2532:
2528:
2525:
2521:
2517:
2510:
2509:
2505:
2502:
2498:
2495:(Dyspnoi?) —
2491:
2490:
2486:
2483:
2479:
2472:
2471:
2467:
2464:
2460:
2456:
2449:
2448:
2444:
2443:
2437:
2435:
2430:
2428:
2424:
2422:
2421:Nemastomoides
2418:
2417:Coal Measures
2413:
2411:
2407:
2403:
2401:
2397:
2393:
2389:
2388:Tetrophthalmi
2385:
2381:
2377:
2373:
2363:
2361:
2357:
2353:
2348:
2346:
2342:
2338:
2337:Cyphophthalmi
2333:
2331:
2326:
2324:
2316:Fossil record
2313:
2311:
2307:
2306:
2301:
2284:
2280:
2275:
2271:
2266:
2262:
2257:
2253:
2250:Martens, 1988
2248:
2244:
2239:
2235:
2234:
2230:
2226:
2219:
2215:
2214:
2210:
2209:Sandokanoidea
2206:
2199:
2195:
2190:
2186:
2183:Thorell, 1889
2181:
2177:
2176:
2172:
2168:
2161:
2160:Phalangodidae
2157:
2156:
2152:
2148:
2141:
2137:
2132:
2128:
2123:
2119:
2114:
2110:
2105:
2104:Nomoclastidae
2101:
2096:
2092:
2087:
2083:
2078:
2074:
2069:
2065:
2060:
2056:
2051:
2047:
2042:
2038:
2033:
2029:
2024:
2020:
2017:Ĺ ilhavĂ˝, 1973
2015:
2014:Agoristenidae
2011:
2010:
2006:
2005:Gonyleptoidea
2002:
1995:
1991:
1986:
1982:
1977:
1973:
1968:
1964:
1963:
1959:
1955:
1948:
1944:
1941:Martens, 2020
1939:
1935:
1930:
1929:Pyramidopidae
1926:
1921:
1917:
1916:
1912:
1908:
1907:
1903:
1899:
1890:
1886:
1883:Forster, 1954
1881:
1877:
1876:
1872:
1868:
1861:
1857:
1852:
1848:
1843:
1839:
1834:
1830:
1829:
1825:
1821:
1820:
1816:
1812:
1811:
1808:Thorell, 1876
1806:
1802:
1793:
1789:
1786:Martens, 1976
1784:
1780:
1775:
1771:
1766:
1762:
1761:
1757:
1753:
1746:
1742:
1737:
1733:
1728:
1724:
1723:
1719:
1715:
1708:
1704:
1703:
1699:
1695:
1694:
1689:
1685:
1676:
1672:
1667:
1666:Protolophidae
1663:
1658:
1654:
1649:
1648:Neopilionidae
1645:
1640:
1636:
1635:
1631:
1630:Phalangioidea
1627:
1620:
1616:
1615:
1611:
1607:
1606:
1601:
1597:
1588:
1587:Parasironidae
1584:
1583:
1581:
1574:
1570:
1565:
1561:
1560:
1556:
1552:
1545:
1541:
1540:
1538:Giribet, 2012
1536:
1532:
1525:
1521:
1520:
1518:Giribet, 2012
1516:
1512:
1505:
1504:Stylocellidae
1501:
1496:
1492:
1491:
1489:Giribet, 2012
1487:
1486:Boreophthalmi
1483:
1482:
1477:
1476:Cyphophthalmi
1473:
1472:
1470:
1465:
1455:
1453:
1449:
1445:
1444:Martin Lister
1441:
1431:
1429:
1425:
1421:
1417:
1413:
1409:
1405:
1402:
1398:
1392:
1382:
1380:
1376:
1375:
1370:
1365:
1363:
1362:South America
1355:
1352:
1349:
1345:
1341:
1337:
1334:
1331:
1329:(1860s–1870s)
1328:
1324:
1321:
1319:(1830s–1850s)
1318:
1314:
1311:
1308:
1305:
1304:
1303:
1300:
1295:
1285:
1283:
1279:
1275:
1274:
1269:
1265:
1254:
1249:
1245:
1241:
1237:
1231:Misconception
1228:
1226:
1222:
1218:
1214:
1210:
1206:
1202:
1198:
1194:
1193:
1188:
1184:
1183:
1178:
1177:
1172:
1170:
1166:
1162:
1158:
1156:
1152:
1148:
1144:
1139:
1137:
1133:
1129:
1125:
1124:
1119:
1115:
1105:
1103:
1099:
1097:
1092:
1088:
1084:
1080:
1076:
1072:
1071:Cyphophthalmi
1068:
1058:
1049:
1047:
1043:
1033:
1031:
1021:
1017:
1013:
1006:
1005:
1000:
991:
989:
984:
974:
965:
963:
959:
948:
946:
942:
938:
934:
930:
920:
918:
914:
910:
906:
901:
892:
890:
886:
882:
881:cyanobacteria
879:
875:
871:
867:
866:
861:
850:
848:
844:
840:
830:
821:
818:
816:
812:
808:
802:
800:
796:
791:
789:
784:
783:Maternal care
780:
779:Paternal care
776:
770:
767:
763:
762:spermatophore
759:
755:
751:
746:
744:
739:
735:
731:
727:
723:
714:
708:
704:
700:
693:
689:
688:
682:
675:
674:
669:
662:
657:
648:
646:
642:
641:
635:
633:
629:
626:is direct as
625:
621:
617:
613:
609:
605:
601:
597:
593:
586:
582:
577:
573:
571:
567:
563:
559:
558:
553:
549:
545:
544:
539:
535:
531:
530:
523:
521:
517:
512:
510:
506:
502:
498:
494:
489:
487:
483:
479:
475:
471:
468:10-segmented
467:
463:
459:
458:cephalothorax
456:
452:
429:
428:
422:
415:
414:
408:
403:
393:
390:
386:
382:
377:
375:
374:cephalothorax
371:
367:
362:
357:
354:
353:Rhynie cherts
349:
347:
342:
340:
339:Tetrophthalmi
336:
332:
328:
324:
323:Cyphophthalmi
320:
316:
312:
308:
304:
300:
296:
292:
288:
277:
274:
270:
265:
264:Tetrophthalmi
261:
259:
256:
254:
251:
249:
246:
244:
243:Cyphophthalmi
241:
240:
238:
233:
228:
223:
220:
217:
216:
213:
210:
207:
206:
203:
200:
197:
196:
193:
190:
187:
186:
183:
180:
177:
176:
173:
170:
167:
166:
161:
156:
152:
148:
147:
142:
138:
133:
128:
124:
116:
111:
106:
101:
96:
91:
86:
81:
76:
71:
66:
61:
55:
44:
41:
37:
33:
19:
6536:
6425:
6419:
6409:
6397:. Retrieved
6391:
6381:
6358:
6352:
6307:
6303:
6293:
6251:(2): e4094.
6248:
6244:
6234:
6191:
6187:
6177:
6165:
6132:
6128:
6121:
6094:
6090:
6080:
6072:
6067:
6059:
6055:
6051:
6040:. Retrieved
6033:the original
5996:
5990:
5965:Science News
5963:
5953:
5934:
5914:
5907:
5897:
5888:
5878:
5867:
5858:
5849:
5815:
5806:
5777:(pp. 83–84).
5774:
5769:
5748:
5738:
5729:
5704:
5695:
5674:
5665:
5631:
5622:
5613:
5604:
5548:
5545:Chemoecology
5544:
5496:
5492:
5482:
5473:
5440:
5436:
5426:
5417:
5396:
5377:
5299:
5295:
5266:
5262:
5252:
5233:
5227:
5194:
5190:
5156:
5152:
5114:
5110:
5067:(2): 85–92.
5064:
5060:
5054:
5021:
5017:
5011:
4987:(1): 58–79.
4984:
4980:
4970:
4937:
4933:
4923:
4890:
4886:
4880:
4855:
4851:
4847:
4841:
4821:
4814:
4803:. Retrieved
4799:the original
4789:
4764:
4760:
4734:. Retrieved
4727:the original
4706:
4700:
4687:
4652:
4648:
4638:
4627:. Retrieved
4623:
4613:
4594:
4587:
4544:
4540:
4529:
4494:
4490:
4480:
4445:
4441:
4391:
4387:
4349:
4317:. Retrieved
4313:the original
4308:
4299:
4288:. Retrieved
4284:
4274:
4263:. Retrieved
4259:
4250:
4238:
4187:Escadabiidae
4169:Sandokanidae
4105:Stygnopsidae
4095:Prostygnidae
4080:Metasarcidae
4075:Manaosbiidae
4070:Gonyleptidae
4045:Askawachidae
4007:Petrobunidae
3997:Beloniscidae
3921:Lomanellidae
3875:Travunioidea
3866:Insidiatores
3829:Phalangiidae
3819:Globipedidae
3570:
3421:Mesostigmata
3290:
3208:Haplocnemata
3190:(harvestmen)
3187:
3181:Stomothecata
3043:
3029:
3019:
3013:
3002:
2988:
2974:
2951:
2934:
2921:
2913:denticulatum
2908:
2896:tuberculatum
2891:
2874:
2863:
2850:
2845:Phalangiidae
2832:
2828:Baltic amber
2812:
2798:
2786:
2769:
2744:
2735:
2704:
2692:
2687:
2684:(Cenomanian)
2668:
2665:(Cenomanian)
2649:
2646:(Cenomanian)
2630:
2627:(Cenomanian)
2611:
2597:(Dyspnoi: â€
2589:
2582:(Dyspnoi: â€
2574:
2559:
2544:
2529:
2516:Eotrogulidae
2514:(Dyspnoi: â€
2506:
2493:Dunlop, 2004
2487:
2482:East Kirkton
2476:(Eupnoi?) —
2474:Dunlop, 2005
2468:
2451:Dunlop, 2004
2445:
2431:
2426:
2425:
2420:
2414:
2405:
2404:
2395:
2391:
2383:
2371:
2369:
2349:
2334:
2327:
2319:
2303:
2297:
2238:Escadabiidae
2227:Superfamily
2218:Sandokanidae
2207:Superfamily
2201:Roewer, 1923
2169:Superfamily
2149:Superfamily
2140:Stygnopsidae
2125:Roewer, 1913
2122:Prostygnidae
2116:Acosta, 2019
2107:Roewer, 1943
2095:Metasarcidae
2089:Roewer, 1943
2086:Manaosbiidae
2077:Gonyleptidae
2053:Roewer, 1913
2032:Askawachidae
2003:Superfamily
1988:Roewer, 1912
1976:Petrobunidae
1956:Superfamily
1950:Roewer, 1912
1909:Superfamily
1869:Superfamily
1854:Briggs, 1971
1824:Travunioidea
1822:Superfamily
1815:Insidiatores
1754:Superfamily
1739:Dresco, 1970
1716:Superfamily
1710:Roewer, 1923
1701:Roewer, 1923
1696:Superfamily
1657:Phalangiidae
1639:Globipedidae
1628:Superfamily
1608:Superfamily
1553:Superfamily
1467:
1452:pre-Linnaean
1447:
1437:
1423:
1404:Rhynie chert
1394:
1372:
1366:
1359:
1333:Eugène Simon
1291:
1271:
1264:urban legend
1261:
1220:
1216:
1212:
1208:
1204:
1200:
1196:
1190:
1180:
1174:
1173:
1171:government.
1160:
1159:
1154:
1150:
1142:
1140:
1135:
1131:
1127:
1121:
1111:
1102:Bothriuridae
1094:
1090:
1082:
1078:
1075:Insidiatores
1074:
1070:
1064:
1055:
1042:stridulating
1039:
1036:Stridulation
1027:
1018:
1014:
1010:
1002:
980:
971:
954:
941:stridulation
926:
902:
898:
870:microhabitat
863:
856:
836:
827:
819:
803:
792:
771:
747:
719:
685:
671:
638:
636:
610:. A pair of
592:scent glands
589:
580:
555:
548:Gonyleptidae
541:
527:
524:
513:
490:
448:
425:
411:
378:
361:phylogenetic
358:
350:
343:
314:
310:
306:
302:
299:colloquially
286:
284:
221:
144:
40:
6703:iNaturalist
6561:Wikispecies
6135:(1): 1–46.
5153:J. Arachnol
5117:: 127–133.
5111:Anim. Behav
4305:"Opiliones"
4256:"Opiliones"
4202:Icaleptidae
4197:Guasiniidae
4179:Zalmoxoidea
4065:Gerdesiidae
3989:Epedanoidea
3974:Suthepiidae
3956:Assamioidea
3947:Grassatores
3898:Travuniidae
3745:Troguloidea
3730:Sabaconidae
3646:Neogoveidae
3408:Holothyrida
3372:Acariformes
3328:Acaromorpha
3299:Schizomida
3198:(scorpions)
3196:Scorpiones
3158:Palpigradi
3125:Chelicerata
3123:Subphylum:
2946:Sabaconidae
2869:Menge, 1854
2817:dentipalpis
2753:Haupt, 1956
2718:Lagerstätte
2569:Mazon Creek
2554:Mazon Creek
2484:, Scotland)
2345:basal clade
2298:The family
2265:Icaleptidae
2256:Guasiniidae
2229:Zalmoxoidea
2163:Simon, 1879
2154:Simon, 1879
2134:Simon, 1879
2068:Gerdesiidae
1958:Epedanoidea
1938:Suthepiidae
1911:Assamioidea
1902:Grassatores
1900:Infraorder
1860:Travuniidae
1836:HadĹľi, 1935
1818:Loman, 1900
1813:Infraorder
1777:Simon, 1872
1768:Simon, 1879
1756:Troguloidea
1736:Sabaconidae
1730:Simon, 1879
1721:Simon, 1879
1678:Simon, 1879
1669:Banks, 1893
1622:Banks, 1892
1613:Banks, 1892
1576:Shear, 1980
1567:Shear, 1980
1564:Neogoveidae
1558:Shear, 1980
1547:Shear, 1993
1533:Infraorder
1527:Shear, 1980
1513:Infraorder
1498:Simon, 1879
1484:Infraorder
1479:Simon, 1879
1458:Systematics
1114:troglobitic
1098:bonariensis
1079:Grassatores
1052:Retaliation
945:retaliation
939:, fleeing,
935:, bobbing,
891:predators.
874:leaf litter
843:aposematism
732:. Some are
673:Protolophus
566:sea spiders
552:Guasiniidae
534:Grassatores
474:opisthosoma
396:Description
202:Chelicerata
198:Subphylum:
52:410–0
6845:Harvestmen
6839:Categories
6188:Cladistics
6042:2012-12-21
5551:(1): 5–7.
5443:(1): 173.
4805:2015-05-29
4736:2007-08-27
4629:2023-07-16
4319:2024-07-10
4290:2017-11-29
4265:2024-07-10
4243:References
4212:Zalmoxidae
4050:Cosmetidae
4017:Tithaeidae
4012:Podoctidae
4002:Epedanidae
3964:Assamiidae
3944:Infraorder
3863:Infraorder
3855:Laniatores
3768:Trogulidae
3665:Infraorder
3635:Infraorder
3626:Pettalidae
3615:Infraorder
3590:Infraorder
3553:Arthropoda
3340:Ricinulei
3277:Amblypygi
3223:Solifugae
3188:Opiliones
3119:Arthropoda
3058:Quaternary
3045:Phalangium
3031:Pellobunus
2944:(Dyspnoi:
2901:(Dyspnoi:
2757:Trogulidae
2755:(Dyspnoi:
2376:ovipositor
2341:Laniatores
2283:Zalmoxidae
2098:Kury, 1994
2062:Kury, 2014
2044:Koch, 1839
2041:Cosmetidae
2026:Kury, 2003
1994:Tithaeidae
1985:Podoctidae
1967:Epedanidae
1920:Assamiidae
1905:Kury, 2002
1805:Laniatores
1792:Trogulidae
1555:Ogoveoidea
1524:Pettalidae
1282:chelicerae
1225:California
1096:Bothriurus
1083:Palpatores
962:Laniatores
951:Thanatosis
929:thanatosis
905:translucid
885:liverworts
860:camouflage
815:Gregarious
807:Laniatores
743:gregarines
734:scavengers
722:omnivorous
707:harvestman
692:tarsomeres
645:Trogulidae
624:copulation
604:book lungs
516:stomotheca
486:chelicerae
464:, and the
387:, and the
385:Tipuloidea
381:arthropods
346:Antarctica
335:Laniatores
307:harvesters
303:harvestmen
258:Laniatores
235:Suborders
192:Arthropoda
18:Harvestmen
6581:Opiliones
6567:Opiliones
6537:Opiliones
6514:(fossils)
6486:Opiliones
6474:Opiliones
6399:April 16,
6326:1664-8021
6267:1314-2836
6210:1096-0031
6149:1096-3642
6113:1095-8312
5883:Research.
5573:0937-7409
5513:0161-8202
5457:0161-8202
5312:CiteSeerX
5003:2177-6199
4852:Behaviour
4649:Curr Biol
4561:0737-4038
4513:1063-5157
4207:Kimulidae
4141:Biantidae
4133:Samooidea
4100:Stygnidae
4055:Cranaidae
4040:Ampycidae
3793:Caddoidea
3735:Taracidae
3651:Ogoveidae
3601:Sironidae
3571:Opiliones
3559:Arachnida
3545:Kingdom:
3481:Haptopoda
3265:Pedipalpi
3254:(spiders)
3172:Dromopoda
3100:Arachnida
2997:Sironidae
2969:Oligocene
2959:(Eupnoi:
2923:Nemastoma
2910:Mitostoma
2884:(Eupnoi:
2876:Leiobunum
2843:(Eupnoi:
2822:(Eupnoi:
2793:Kimulidae
2761:Geiseltal
2722:Koonwarra
2541:, France)
2520:Commentry
2499:(Western
2410:Edinburgh
2392:E. sheari
2366:Paleozoic
2360:Paleozoic
2274:Kimulidae
2180:Biantidae
2171:Samooidea
2131:Stygnidae
2050:Cranaidae
2023:Ampycidae
1803:Suborder
1745:Taracidae
1686:Suborder
1610:Caddoidea
1598:Suborder
1573:Ogoveidae
1495:Sironidae
1474:Suborder
1434:Etymology
1420:Dromopoda
1416:solifuges
1408:scorpions
1397:arachnids
1385:Phylogeny
1278:crane fly
1248:Solifugae
1030:Leiobunum
988:Pholcidae
909:carapaces
865:Leiobunum
795:nocturnal
758:fossorial
748:Although
612:spiracles
520:pedipalps
466:posterior
301:known as
295:arachnids
287:Opiliones
273:Diversity
227:Sundevall
222:Opiliones
212:Arachnida
178:Kingdom:
172:Eukaryota
46:Opiliones
6612:BugGuide
6588:BioLib:
6546:Wikidata
6452:16024358
6344:25120562
6285:25425936
6226:85408627
6218:34772266
6157:84029705
6021:19495718
5943:Archived
5599:300–308.
5581:31558441
5521:83994309
5465:85822815
5342:10906916
5334:16273426
5211:26470886
5173:85621336
5131:54254258
5089:83850344
5046:85961182
4962:85846440
4954:40233818
4915:18019589
4872:49329266
4781:38401545
4723:85918640
4679:24726154
4579:35137183
4521:30917194
4472:21863011
4418:24726154
4146:Samoidae
3852:Suborder
3801:Caddidae
3781:Suborder
3687:Suborder
3667:(Indet.)
3579:Suborder
3551:Phylum:
3547:Animalia
3413:Ixodida
3307:Uropygi
3252:Araneae
3117:Phylum:
3113:Animalia
3111:Kingdom
3020:A. lawei
2939:claviger
2926:incertum
2879:longipes
2824:Caddidae
2803:silhavyi
2778:Samoidae
2749:longipes
2746:Trogulus
2732:Cenozoic
2726:Victoria
2707:Triassic
2606:Mesozoic
2501:Missouri
2463:Scotland
2396:H. argus
2380:tracheae
2356:Mesozoic
2352:Cenozoic
2330:Silurian
2189:Samoidae
1619:Caddidae
1401:Devonian
1288:Research
1276:and the
1268:venomous
1087:Ctenidae
1067:ozopores
1061:Chemical
994:Autotomy
968:Freezing
937:autotomy
933:freezing
651:Behavior
620:gonopore
608:tracheae
600:quinones
596:ozopores
509:antennae
505:medially
501:carapace
455:anterior
188:Phylum:
182:Animalia
168:Domain:
127:Holocene
123:Devonian
6822:1300999
6443:1599874
6335:4112917
6310:: 255.
6276:4238074
6075:, p.321
6029:9570512
6001:Bibcode
5553:Bibcode
5304:Bibcode
5219:1994257
5197:: 1–7.
5069:Bibcode
5026:Bibcode
4895:Bibcode
4657:Bibcode
4570:8845124
4450:Bibcode
4448:: 444.
4396:Bibcode
3690:Dyspnoi
3557:Class:
3471:Extinct
3415:(ticks)
3034:proavus
2967:, USA (
2936:Sabacon
2837:ramiger
2765:Germany
2678:Myanmar
2659:Myanmar
2640:Myanmar
2621:Myanmar
2571:, U.S.)
2556:, U.S.)
2503:, U.S.)
2434:Permian
2281:Family
2272:Family
2263:Family
2254:Family
2245:Family
2236:Family
2216:Family
2196:Family
2187:Family
2178:Family
2158:Family
2138:Family
2129:Family
2120:Family
2111:Family
2102:Family
2093:Family
2084:Family
2075:Family
2066:Family
2057:Family
2048:Family
2039:Family
2030:Family
2021:Family
2012:Family
1992:Family
1983:Family
1974:Family
1965:Family
1945:Family
1936:Family
1927:Family
1918:Family
1887:Family
1878:Family
1858:Family
1849:Family
1840:Family
1831:Family
1790:Family
1781:Family
1772:Family
1763:Family
1743:Family
1734:Family
1725:Family
1705:Family
1688:Dyspnoi
1673:Family
1664:Family
1655:Family
1646:Family
1637:Family
1617:Family
1585:Family
1571:Family
1562:Family
1542:Family
1522:Family
1502:Family
1493:Family
1323:L. Koch
1246:in the
1240:Chelate
1169:Spanish
1024:Fleeing
977:Bobbing
958:Dyspnoi
889:diurnal
878:epizoic
853:Crypsis
847:mimicry
839:crypsis
799:diurnal
788:instars
726:insects
684:A male
585:spiders
540:nests,
538:termite
536:) from
470:abdomen
462:prosoma
451:tagmata
370:Araneae
368:(order
366:spiders
331:Dyspnoi
253:Dyspnoi
218:Order:
208:Class:
6796:133884
6767:NZOR:
6656:1OPIOO
6552:Q19116
6450:
6440:
6369:
6342:
6332:
6324:
6283:
6273:
6265:
6224:
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5648:87–99.
5579:
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4567:
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4470:
4416:
4360:
3784:Eupnoi
3568:Order
3139:Extant
3102:orders
2855:ovalis
2852:Opilio
2788:Kimula
2738:Eocene
2524:France
2459:Rhynie
2400:Eupnoi
1600:Eupnoi
1448:opilio
1414:, and
1244:chelae
1219:, and
1118:Brazil
983:Eupnoi
845:, and
811:Eupnoi
766:mating
570:Eupnoi
497:scutum
493:dorsal
453:, the
430:) body
359:Their
337:, and
327:Eupnoi
319:extant
248:Eupnoi
229:, 1833
121:Early
6817:WoRMS
6804:Plazi
6760:43271
6734:82753
6721:10452
6716:IRMNG
6708:47367
6669:15591
6604:24643
6591:16797
6222:S2CID
6153:S2CID
6036:(PDF)
6025:S2CID
5987:(PDF)
5577:S2CID
5517:S2CID
5461:S2CID
5338:S2CID
5215:S2CID
5169:S2CID
5127:S2CID
5085:S2CID
5042:S2CID
4958:S2CID
4950:JSTOR
4911:S2CID
4868:S2CID
4730:(PDF)
4719:S2CID
4697:(PDF)
4547:(2).
3354:Acari
3054:Italy
2814:Caddo
2780:?) —
2670:indet
2323:amber
1187:Texas
738:fecal
730:fungi
703:Mites
661:skink
616:tibia
482:venom
313:, or
291:order
6783:4428
6755:NCBI
6729:ITIS
6690:GBIF
6651:EPPO
6617:2405
6599:BOLD
6448:PMID
6401:2014
6367:ISBN
6340:PMID
6322:ISSN
6281:PMID
6263:ISSN
6214:PMID
6206:ISSN
6145:ISSN
6109:ISSN
6017:PMID
5920:ISBN
5569:ISSN
5509:ISSN
5453:ISSN
5382:ISBN
5330:PMID
5238:ISBN
5207:PMID
4999:ISSN
4827:ISBN
4777:PMID
4675:PMID
4600:ISBN
4575:PMID
4557:ISSN
4517:PMID
4509:ISSN
4468:PMID
4414:PMID
4358:ISBN
3291:s.l.
3050:Rome
2963:) —
2826:) —
2759:) —
2394:and
1179:and
1134:and
1112:All
960:and
883:and
663:tail
632:eggs
564:and
478:oval
285:The
60:Preęž’
6742:NBN
6695:907
6643:249
6638:EoL
6630:3Q2
6625:CoL
6576:AFD
6438:PMC
6430:doi
6426:272
6330:PMC
6312:doi
6271:PMC
6253:doi
6196:doi
6137:doi
6133:174
6099:doi
6095:105
6009:doi
5561:doi
5501:doi
5445:doi
5322:doi
5271:doi
5199:doi
5195:121
5161:doi
5119:doi
5077:doi
5034:doi
4989:doi
4942:doi
4903:doi
4860:doi
4856:151
4769:doi
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4665:doi
4565:PMC
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