69-590: The Pettalidae are a family of harvestmen with 75 described species in 10 genera . Several undescribed species are known or assumed in some genera. Pettalus is a name from Greek mythology that appears in Ovid 's Metamorphoses . All species except the cave-dwelling South African Speleosiro argasiformis spend their entire life cycle in leaf litter . They are two to five millimeters long, usually with an oval shaped body. Although all Pettalidae except Parapurcellia have eyes, these were long thought to be absent in
138-451: A pre-Linnaean 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. 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
207-409: A 2012 phylogenetic analysis, a new Cyphophthalmi taxonomy was proposed. The Cyphophthalmi are currently represented by the following taxa, which represent approximately 200 species, although there are a significant number of undescribed species that have been collected, and the current number of species is believed to be a severe under-representation. The monophyly of most subgroups of Cyphophthalmi
276-618: 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 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 gregarines . Although parthenogenetic species do occur, most harvestmen reproduce sexually . Except from small fossorial species in
345-407: A distance. 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 tagmata , the anterior cephalothorax or prosoma , and the posterior 10-segmented abdomen or opisthosoma . The most easily discernible difference between harvestmen and spiders is that in harvestmen,
414-456: 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. 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
483-732: A long period. Indeed, one species discovered in China, Mesobunus martensi , 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 Sclerosomatidae . The Swedish naturalist and arachnologist Carl Jakob Sundevall (1801–1875) honored the naturalist Martin Lister (1638–1712) by adopting Lister's term Opiliones for this order, known in Lister's days as "harvest spiders" or "shepherd spiders", from Latin opilio , "shepherd"; Lister characterized three species from England (although not formally describing them, being
552-459: A modern genus, Siro platypedibus Dunlop & Giribet, 2003 (Sironidae), was described from Paleogene aged Bitterfeld amber of eastern Germany . Another of this genus was described from Eocene Baltic amber as Siro balticus Dunlop & Mitov, 2011 . A third fossil, Palaeosiro burmanicum Poinar, 2008 , has been described from the Cretaceous ( Cenomanian ) aged Burmese amber . It
621-428: A pair median eyes also have two sets of vestigial eyes: one median pair (homologous to those of horseshoe crabs and sea spiders ), 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
690-418: A predator after direct or indirect contact, including thanatosis , freezing , bobbing, autotomy , fleeing, stridulation , retaliation and chemical secretions. 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
759-611: A predator. 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 spiders by causing injury, allowing the harvestman to escape from predation. Harvestmen are well known for being chemically protected. They exude strongly odored secretions from their scent glands, called ozopores , that act as
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#1732886267823828-510: A scientifically neglected group. Description of new taxa has always been dependent on the activity of a few dedicated taxonomists. Carl Friedrich Roewer described about a third (2,260) of today's known species from the 1910s to the 1950s, and published the landmark systematic work Die Weberknechte der Erde (Harvestmen of the World) in 1923, with descriptions of all species known to that time. Other important taxonomists in this field include: Since
897-516: A shield against predators; this is the most effective defense they use which creates a strong and unpleasant taste. In Cyphophthalmi the scent glands release naphthoquinones, chloro-naphthoquinones and aliphatic methyl ketones, Insidiatores use nitrogen-containing substances, terpenes, aliphatic ketones, and phenolics, while Grassatores use alkylated phenolics and benzoquinones, and Palpatores use substances like naphthoquinones, methyl- and ethyl-ketones. These secretions have successfully protected
966-473: A structure which is not inserted inside the female, but used to deposit a spermatophore. In addition, they are the only harvestmen to lack a genital operculum, and instead exhibit a completely open gonopore. Mating in Cyphophthalmi is not well studied, but in some species at least, males and females do engage in direct contact. They further differ from most harvestmen in that the first 8 opisthosomal tergites and
1035-492: A variety of primary and secondary defences against predation, ranging from morphological traits such as body armour to behavioral responses to chemical secretions. Some of these defences have been attributed and restricted to specific groups of harvestmen. Primary defences help the harvestmen avoid encountering a potential predator and include crypsis , aposematism , and mimicry . Particular patterns or colour markings on harvestmen's bodies can reduce detection by disrupting
1104-403: A variety of shapes, depending on species. The function of the adenostyle is currently unknown, but is presumably associated with chemical glands. Aside from the adenostyle, males in many species possess glandular pores on either the underside of the opisthosoma or the anal region, although these glands are not present in all species. Males in several families also possess structural modifications of
1173-507: Is monophyletic , although it is at the moment (2007) unclear what the nearest relatives are. It probably originated in the southern part of Gondwana. Parsimony analysis suggests it could be a sister group to the remaining Cyphophthalmi , though this could also be the case for the Stylocellidae , or it could be related to the Sironidae , or specifically to the sironid genus Suzukielus . It
1242-558: Is a compilation of interfamilial relationships recovered from several recent phylogenetic studies, although the placement and even monophyly of several taxa are still in question. The family Stygophalangiidae (one species, Stygophalangium karamani ) from underground waters in North Macedonia is sometimes misplaced in the Phalangioidea. It is not a harvestman. Despite their long history, few harvestman fossils are known. This
1311-485: Is also used for two other distantly related groups of arthropods , the crane flies of the superfamily Tipuloidea , and the cellar spiders 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 better observe their wandering flocks from
1380-514: Is believed that they did not travel between separate landmasses, they make an interesting subject for biogeography . Each of the six currently recognized families has a distinct distribution: Colombia shows the largest diversity of cyphophthalmids among countries in South America. This could reflect the large number of ecosystems found there, but could also be due to a high rate of sampling in this country. A fossil cyphophthalmid assigned to
1449-432: Is formed by extensions of the coxae of the pedipalps and the first pair of legs. 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 was reported in 2014. This find suggested that
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#17328862678231518-510: Is in rainforest patches in northwest Argentina which are in an area being dramatically destroyed by humans. The cave-living Picunchenops spelaeus 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. Maiorerus randoi has only been found in one cave in the Canary Islands . It is included in
1587-529: Is known about their behavior, though they likely subsist mostly by scavenging and preying on minute arthropods. They have low dispersal rates and consequently high endemism . Cyphophthalmi differ from other harvestmen in a number of key ways. Like all harvestmen, they have a pair of ozopores located on the sides of the prosoma. Unlike other harvestmen, however, the ozopores in Cyphophthalmi are located on elevated cone-shaped structures known as ozophores. Although members of most families are eyeless, most members of
1656-508: 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 amber . 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 Silurian , or even earlier. A recent molecular study of Opiliones, however, dated
1725-404: Is most likely a member of the suborder Eupnoi (true daddy-longlegs). Harvestmen have a pair of prosomatic defensive scent glands ( ozopores ) that secrete a peculiar-smelling fluid when disturbed. In some species, the fluid contains noxious quinones . They do not have book lungs , and breathe through tracheae . A pair of spiracles is located between the base of the fourth pair of legs and
1794-399: Is possible that the Cyphophthalmi dispersed across Australia while the central region was covered with Nothofagus rainforest (until 37 mya), or that the ancestors of the two genera independently dispersed from adjacent landmasses now separate from Australia. Parapurcellia from eastern South Africa is sister to all other Pettalidae, while Purcellia from western South Africa is sister to
1863-490: Is seen with multiple long-legged species in the Leiobunum 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. Multiple species within the Laniatores and Dyspnoi possess stridulating organs, which are used as intraspecific communication and have also been shown to be used as a second line of defense when restrained by
1932-452: 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 their legs, and leg loss reduces locomotion, speed, climbing ability, sensory perception, food detection, and territoriality. Autotomised legs provide
2001-617: Is thought to be commonly used during daylight, when they could be easily seen by any predators. Other harvestmen may exhibit mimicry to resemble other species' appearances. Some Gonyleptidae individuals that produce translucid secretions have orange markings on their carapaces . This may have an aposematic role by mimicking the colouration of glandular emissions of two other quinone-producing species. Mimicry ( Müllerian mimicry ) occurring between Brazilian harvestmen that resemble others could be explained by convergent evolution . Secondary defences allow for harvestmen to escape and survive from
2070-621: Is unrelated to the Troglosironidae that are endemic to New Caledonia . The main lineages of the family may have arisen rapidly, possibly during the rapid expansion of Glossopteris forests that were predominant in temperate Gondwana. Pettalidae were likely present throughout the forests of Antarctica , which formed a land bridge between Australia and South America up until circa 50 million years ago (mya). The Australian genera Austropurcellia (Eastern Australia: Queensland) and Karripurcellia (Western Australia) are not sister groups. It
2139-594: Is used as a second line of defence when detected by a potential predator and is commonly observed within the Dyspnoi and Laniatores suborders, with individuals becoming rigid with legs either retracted or stretched. 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. To deflect attacks and enhance escape, long-legged species – commonly known as daddy long-legs – from
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2208-638: The Cenozoic , three from the Mesozoic , and at least seven from the Paleozoic . Cyphophthalmi Cyphophthalmi is a suborder of harvestmen , colloquially known as mite harvestmen . Cyphophthalmi comprises 36 genera, and more than two hundred described species. The six families are currently grouped into three infraorders: the Boreophthalmi, Scopulophthalmi, and Sternophthalmi. Cyphophthalmi are smaller than
2277-509: The Devonian Rhynie chert , 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 scorpions , pseudoscorpions , and solifuges ; these four orders form the clade Dromopoda . The Opiliones have remained almost unchanged morphologically over
2346-606: The Eupnoi 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 individuals, which are all trembling at the same time. Cellar spiders ( Pholcidae ) that are commonly mistaken for daddy long-legs (Opiliones) also exhibit this behavior when their webs are disturbed or even during courtship. Autotomy
2415-457: The Laniatores , Dyspnoi and Eupnoi , is known as Phalangida . The extinct suborder, Tetrophthalmi , shares several features in common with Cyphophthalmi, and these two suborders may represent sister taxa. They are grouped into the following infraorders: Boreophthalmi, Scopulophthalmi, and Sternophthalmi. The Boreophthalmi and Sternophthalmi together form a monophyletic clade, and comprise 5 of
2484-521: The Laniatores , and more than 70,000 in certain Eupnoi . Gregarious 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. Harvestmen clean their legs after eating by drawing each leg in turn through their jaws. Predators of harvestmen include a variety of animals, including some mammals, amphibians, and other arachnids like spiders and scorpions. Opiliones display
2553-473: The leaf litter found in their environments. Some hard-bodied harvestmen have epizoic cyanobacteria and liverworts growing on their bodies that suggest potential benefits for camouflage against large backgrounds to avoid detection by diurnal predators. 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
2622-573: The scutum , which in most such species is fused with the carapace . 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 medially on the surface to form two plates beside each other. The second pair of legs is longer than the others and function as antennae or feelers. In short-legged species, this may not be obvious. The feeding apparatus ( stomotheca ) differs from most arachnids in that Opiliones can swallow chunks of solid food, not only liquids. The stomotheca
2691-479: The 1990s, study of the biology and ecology of harvestmen has intensified, especially in South America . Early work on the developmental biology of Opiliones from the mid-20th century was resurrected by Prashant P. Sharma , who established Phalangium opilio as a model system for the study of arachnid comparative genomics and evolutionary-developmental biology . Harvestmen are ancient arachnids . Fossils from
2760-708: The 400-million-year-old Rhynie cherts 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. Their phylogenetic 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 spiders (order Araneae ),
2829-533: The 6 families of Cyphphthalmi. The Scopulophthalmi, with only 1 family, the Pettalidae, forms the sister group to all the other families. Within the Sternophthalmi, the families Ogoveidae and Neogoveidae are most closely related, and form the superfamily Ogoveoidea, which is the sister group to the family Troglosironidae. The systematics of the Boreophthalmi, however, is not yet fully resolved. In one recent analysis,
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2898-624: The Boreophthalmi were recovered as paraphyletic with respect to the Sternophthalmi. Even within the Boreophthalmi, the Sironidae has been recovered either as sister family to Stylocellidae, or as paraphyletic with respect to Stylocellidae. In addition, there are 3 genera of Cyphophthalmi with uncertain affinities that have not been placed in any family or infraorder. Pettalidae Sironidae Stylocellidae Neogoveidae Ogoveidae Troglosironidae Pettalidae Stylocellidae Sironidae Neogoveidae Ogoveidae Troglosironidae Originally, Cyphophthalmi
2967-551: The Brazilian national list of endangered species, all of them cave-dwelling: Giupponia chagasi , Iandumoema uai , Pachylospeleus strinatii and Spaeleoleptes spaeleus . Several Opiliones in Argentina appear to be vulnerable, if not endangered. These include Pachyloidellus fulvigranulatus , which is found only on top of Cerro Uritorco , the highest peak in the Sierras Chicas chain (provincia de Cordoba) and Pachyloides borellii
3036-522: The Carboniferous. Another study, in 2014, recovered a diversification age of approximately 340 MYA. A more recent study in 2017, however, recovered the diversification of the modern Cyphophthalmi lineages to have occurred more recently, during the Jurassic, with the emergence of the suborder as a whole at less than 300 MYA. Cyphophthalmi is one of the two major lineages of harvestmen; the other, containing
3105-820: The Catálogo Nacional de especies amenazadas (National catalog of threatened species) from the Spanish government. Texella reddelli and Texella reyesi are listed as endangered species in the United States. Both are from caves in central Texas . Texella cokendolpheri from a cave in central Texas and Calicina minor , Microcina edgewoodensis , Microcina homi , Microcina jungi , Microcina leei , Microcina lumi , and Microcina tiburona from around springs and other restricted habitats of central California are being considered for listing as endangered species, but as yet receive no protection. An urban legend claims that
3174-652: The Chilean Chileogovea . Western South Africa and southern South America were last connected during the Late Jurassic , about 150 mya. Likewise, the monotypic Neopurcellia from New Zealand appears as the sister group to all Pettalidae except for Parapurcellia , instead of being monophyletic with the other two New Zealand genera, which themselves appear as sister groups in Bayesian analysis , but not in direct optimization parsimony analyses. The family Pettalidae contains
3243-532: 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 cephalothorax . 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. English speakers may colloquially refer to species of Opiliones as "daddy longlegs" or "granddaddy longlegs", but this name
3312-406: The abdomen, with one opening on each side. In more active species, spiracles are also found upon the tibia of the legs. They have a gonopore on the ventral cephalothorax, and the copulation is direct as male Opiliones have a penis , unlike other arachnids. All species lay eggs . Typical body length does not exceed 7 mm (0.28 in), and some species are smaller than 1 mm, although
3381-456: The anal plate and males of certain species in the family Stylocellidae possess a modified patch of cuticle near the adenostyle known as the Rambla's organ. The exact function of these glands and structures is currently unknown. Like most harvestmen, Cyphophthalmi rely heavily on chemical secretions for defense. This defensive character is particularly well developed in Cyphophthalmi, when compared to
3450-500: 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 camouflage when they remain motionless. Within the genus Leiobunum are multiple species with cryptic colouration that changes over ontogeny to match the microhabitat used at each life stage. Many species have also been able to camouflage their bodies by covering with secretions and debris from
3519-421: The connection between the cephalothorax and abdomen is broad, so that the body appears to be a single oval structure. Other differences include the fact that Opiliones have no venom glands in their chelicerae and thus pose no danger to humans. 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 dorsal shield called
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#17328862678233588-594: The eyes in Cyphophthalmi are not homologous to the eyes of other harvestmen. Many cave-adapted species are eyeless, such as the Brazilian Caecobunus termitarum ( Grassatores ) from termite nests, Giupponia chagasi ( Gonyleptidae ) from caves, most species of Cyphophthalmi, and all species of the Guasiniidae . However, recent work studying the embryonic development of the species Phalangium opilio and some Laniatores revealed that harvestman in addition to
3657-423: The families Pettalidae and Stylocellidae do have eyes located on or near the ozophores. These eyes, however, are not homologous to the eyes of other harvestmen, and are instead derived from ancestral lateral eyes, instead of ancestral median eyes, as is the case for other harvestmen. The male genitalia of Cyphophthalmi is unique among harvestmen, as males lack a true penis, and instead have a short spermatopositor ,
3726-812: The family, mainly because they cannot be seen by scanning electron microscopy . They are often incorporated at the base of the ozophores and typically lack lenses. The members of this family are distributed throughout former temperate Gondwana , with genera in Chile , South Africa , Madagascar , Sri Lanka , eastern and western Australia , and New Zealand , where they are most diverse by far, with 29 species and subspecies found in three genera. Parapurcellia (eastern South Africa) Neopurcellia (New Zealand) Rakaia (New Zealand) Aoraki (New Zealand) Purcellia (western South Africa) Chileogovea (Chile) Karripurcellia (Western Australia) Pettalus (Sri Lanka) Austropurcellia (Queensland) The family Pettalidae
3795-652: The following genera and species: Harvestmen The Opiliones (formerly Phalangida) are an order of arachnids , colloquially known as harvestmen , harvesters , harvest spiders , or daddy longlegs . As of July 2024 , over 6,650 species of harvestmen have been discovered worldwide, although the total number of extant species may exceed 10,000. The order Opiliones includes five suborders: Cyphophthalmi , Eupnoi , Dyspnoi , Laniatores , and Tetrophthalmi , which were named in 2014. Representatives of each extant suborder can be found on all continents except Antarctica . Well-preserved fossils have been found in
3864-490: The harvestman is the most venomous 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 Pholcus phalangioides and the crane fly , which are both also called a "daddy longlegs"). None of the known species of harvestmen have venom glands; their chelicerae are not hollowed fangs but grasping claws that are typically very small and not strong enough to break human skin. Harvestmen are
3933-565: The harvestmen against wandering spiders ( Ctenidae ), wolf spiders (Lycosidae) and Formica exsectoides ants. However, these chemical irritants are not able to prevent four species of harvestmen being preyed upon by the black scorpion Bothriurus bonariensis ( Bothriuridae ). These secretions contain multiple volatile compounds that vary among individuals and clades. All troglobitic species (of all animal taxa) are considered to be at least threatened in Brazil . Four species of Opiliones are on
4002-568: The largest known species, Trogulus torosus ( Trogulidae ), 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. Many species are omnivorous , eating primarily small insects and all kinds of plant material and fungi . Some are scavengers , feeding upon dead organisms, bird dung, and other fecal material. Such
4071-557: The male is solely responsible for guarding the eggs resulting from multiple partners, often against egg-eating females , and cleaning the eggs regularly. Paternal care has evolved at least three times independently: once in the clade Progonyleptoidellinae + Caelopyginae, once in the Gonyleptinae, and once in the Heteropachylinae. Maternal care in opiliones probably evolved due to natural selection, while paternal care appears to be
4140-435: 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. 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
4209-558: The more familiar "daddy long-legs" harvestmen, with adults ranging from 1 to 7 mm in length. Moreover, their legs are relatively short compared to most other harvestmen, typically shorter than the body. Some superficially resemble mites , which is where they get their common name. Their coloration is almost always some shade of brown, with a heavily sclerotized body, and they are quite inconspicuous, residing in leaf litter or in caves . Many Cyphophthalmi are eyeless, and presumably rely on olfactory cues to find food and mates. Very little
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#17328862678234278-522: The more familiar Eupnoi and Dyspnoi. Cyphophthalmi possess more elaborate musculature associated with the defensive glands than other harvestmen, and the secretions, which primarily consist of a variety of naphthoquinones and ketones, are thought to be more potent than in Eupnoi and Dyspnoi. Cyphophthalmids are found on all continents, with the exception of Antarctica , where they probably also once lived. As they did not disperse onto any oceanic islands, and it
4347-424: The nerves to the muscles to extend the leg and then the leg relaxes between signals. While some harvestman's legs twitch 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. Individuals that are able to detect potential threats can flee rapidly from attack. This
4416-566: The origin of the order at about 473 million years ago (Mya), during the Ordovician. No fossils of the Cyphophthalmi or Laniatores much older than 50 million years are known, despite the former presenting a basal clade , and the latter having probably diverged from the Dyspnoi more than 300 Mya. Naturally, most finds are from comparatively recent times. More than 20 fossil species are known from
4485-437: The prosomal carapace are fused together in what is known as a scutum completum , although this feature is also known from the unrelated family Sandokanidae . Like most animals, Cyphophthalmi express morphological distinctions between male and female individuals. Male Cyphophthalmi possess a structure on their fourth pair of tarsi known as an adenostyle. The adenostyle usually appears as a small hornlike projection, but can take
4554-746: 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 four to eight nymphal instars to reach maturity, with most known species having six instars. Most species are nocturnal and colored in hues of brown, although a number of diurnal species are known, some of which have vivid patterns in yellow, green, and black with varied reddish and blackish mottling and reticulation. 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
4623-437: The suborder Cyphophthalmi, where the males deposit a spermatophore , mating 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 guards the female after copulation, and in many species,
4692-416: Was also assigned to Sironidae but is now thought to belong to Stylocellidae (a more typical Asian family). As the fossil record for Cyphophthalmi is so sparse, recent studies have attempted to predict the diversification time of the suborder by using molecular clocks and comparing gene sequences among living taxa. One 2012 study estimated the earliest diversification of the suborder at approximately 332 MYA, in
4761-485: Was recognized as a single family consisting of two subfamilies, Sironini and Stylocellini. This classification scheme existed until 1980, when a new taxonomy was proposed, separating 5 families into the now-defunct infraorders Temperophthalmi (consisting of Petallidae, Sironidae and, later, Troglosironidae) and Tropicophthalmi (consisting of Stylocellidae, Ogoveidae and Neogoveidae). These two infraorders were never recovered as monophyletic in subsequent studies, and, following
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