Opiliones

For other arthropods called "daddy longlegs", see Daddy longlegs.
Opiliones
Temporal range: 400–0 Ma

Early Devonian – Holocene

Opiliones (Hadrobunus grandis) showing its body structure and long legs. A sole pair of eyes and unitary cephalothorax differentiate Opiliones from other members of the class Arachnida
Scientific classification
Kingdom: Animalia
Phylum: Arthropoda
Class: Arachnida
Subclass: Dromopoda
Order: Opiliones
Sundevall, 1833
Suborders
Diversity
5 suborders, > 6,500 species

Opiliones (/ˈoʊpɪliːˈoʊniːs/; formerly Phalangida) is an order of arachnids commonly known as harvestmen. As of December 2011, over 6,500 species of harvestmen have been discovered worldwide,[1] although the total number of extant species may exceed 10,000.[2] The order Opiliones includes five suborders: Cyphophthalmi, Eupnoi, Dyspnoi, Laniatores, and the recently named Tetrophthalmi.[3] With the exception of Dyspnoi (which is restricted to North America and Eurasia), representatives of each can be found on every continent except Antarctica. Well-preserved fossils have been found in the 400-million-year-old Rhynie cherts of Scotland, and 305-million-year-old rocks in France, which look surprisingly modern, indicating that their basic body plan appeared very early on,[4] and, at least in some taxa, has changed little since that time. Their phylogenetic position within Arachnida is disputed: their closest relatives may be the mites (Acari) or the Novogenuata (the Scorpiones, Pseudoscorpiones and Solifugae).[5] Although superficially similar to and often confused with spiders (order Araneae), Opiliones is a distinct order that is not closely related to spiders within Arachnida. They can be easily distinguished from even long-legged spiders by their fused body regions and single pair of eyes in the middle of their cephalothorax (spiders have an 'abdomen' that is separated from the cephalothorax by a constriction, as well as three to four pairs of eyes, usually around the margins of their cephalothorax).

English speakers colloquially refer to species of Opiliones as "daddy longlegs" or "granddaddy longlegs", but this name is also used for two other unrelated groups of arthropods: the crane flies of the family Tipulidae, and the cellar spiders of Pholcidae, most likely because of their similar appearance. They 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 a distance.[6]

Physical description

Main article: Harvestman anatomy
Tropical harvestman (Pachyloidellus goliath)
North European harvestman (Leiobunum rotundum) body

Opiliones are known for having exceptionally long legs relative to their body size; however, some species are short-legged. As in the Araneae, the body in the Opiliones has two tagmata, the anterior cephalothorax or prosoma, and the posterior ten-segmented abdomen or opisthosoma. The most obvious 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 oval structure. Other differences are that Opiliones have no venom glands in their chelicerae and therefore 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 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 divided medially on the surface to form two plates beside each other. The second pair of legs are longer than the others and they 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 is formed by extensions from the pedipalps and the first pair of legs.

Opiliones have a single pair of eyes in the middle of the head, oriented sideways. However, there are eyeless species, such as the Brazilian Caecobunus termitarum (Grassatores) from termite nests, Giupponia chagasi (Gonyleptidae) from caves, and all species of Guasiniidae.[7]

A harvestman (a male Phalangium opilio), showing the almost fused arrangement of abdomen and cephalothorax that distinguishes these arachnids from spiders.

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. Between the base of the fourth pair of legs and the abdomen a pair of spiracles are located, 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.

The legs continue to twitch after they are detached. This is because there are 'pacemakers' 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 will twitch for a minute, other kinds have been recorded to twitch for up to an hour. The twitching has been hypothesized as a means to keep the attention of a predator while the harvestman escapes.[2]

Typical body length does not exceed 7 millimetres (0.28 in), and some species are smaller than one mm, although the largest known species Trogulus torosus (Trogulidae) grow as long as 22 millimetres (0.87 in).[2] The leg span of many species is much greater than the body length and sometimes exceeds 160 millimetres (6.3 in) all the way up to 340 millimetres (13 in) in south eastern Asia.[8] Most species live for a year.

Behavior

Harvestman eating a skink tail
Male harvestman Opilio canestrinii cleaning its legs
A male Phalangium opilio, showing the long legs and the tarsomeres (the many small segments making up the end of each leg)
Mites parasitising a harvestman
Gregarious behaviour in Opiliones

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 broad range is unusual in other 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.[2]

Although parthenogenetic species do occur, most harvestmen reproduce sexually. Mating involves direct copulation, rather than the deposition of a spermatophore. The males of some species offer a secretion from their chelicerae to the female before copulation. Sometimes the male guards the female after copulation and, in many species, the males defend territories. The females lay eggs shortly after mating or anytime up to several months later. Some species build nests for this purpose. A unique feature of harvestmen is that in some species the male is solely responsible for guarding the eggs resulting from multiple partners, often against egg-eating females, and subjecting the eggs to regular cleaning. 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.[2]

Most species are nocturnal and colored in hues of brown, although there are a number of diurnal species, some of which have vivid patterns in yellow, green and black with varied reddish and blackish mottling and reticulation.

To deal with predators such as birds, mammals, amphibians and spiders, some species glue debris onto their body, while many play dead when disturbed. Many species practice autotomy; they detach their legs, which keep on moving for a period of time after, presumably to distract predators. Especially long-legged species vibrate their body ("bobbing"), probably also to confuse predators. This is similar to the behavior of the similar looking but unrelated cellar spider, which vibrates wildly in its web when touched. Scent glands emit substances that can deter larger predators, but are also effective against ants.[2]

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 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 of being eaten.[2]

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

Endangered status

All troglobitic species (of all animal taxa) are considered to be at least threatened in Brazil. There are four species of Opiliones in the Brazilian National List for endangered species, all of them cave-dwelling species. 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 is in rainforest patches in North West 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 and therefore they receive no protection.

Maiorerus randoi has only been found in one cave in the Canary Islands. It is included in 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.

Misconception

Chelate (pincer-like) chelicerae typical of harvestmen (200x magnification); these chelicerae are homologous to chelicerae that take the form of fangs in spiders or chelae in the Solifugae.

An urban legend claims that the harvestman is the most venomous animal in the world,[9] but possesses fangs too short or a mouth too round and small to bite a human and therefore is not dangerous (the same myth applies to Pholcus phalangioides and the cranefly, which are both also called a 'daddy longlegs').[10] This is untrue on several counts. None of the known species of harvestmen has venom glands; their chelicerae are not hollowed fangs but grasping claws that are typically very small and not strong enough to break human skin.

Research

Harvestmen are 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: Pierre Latreille (18th century) Carl Ludwig Koch, Maximilian Perty (1830s-1850s) L. Koch, Tord Tamerlan Teodor Thorell (1860s-1870s) Eugène Simon, William Sørensen (1880s-1890s) James C. Cokendolpher, Raymond Forster, Jürgen Gruber, Reginald Frederick Lawrence, Jochen Martens, Cândido Firmino de Mello-Leitão (20th century) Gonzalo Giribet, Adriano Brilhante Kury, Tone Novak (21st century).

Since the 1990s, study of the biology and ecology of harvestmen has intensified, especially in South America.[2]

Phylogeny

Main article: Harvestman phylogeny

Harvestmen are very old arachnids. Fossils from the Devonian Rhynie chert, 410 million years ago, already show characteristics like tracheae and sexual organs, proving that the group has lived on land since that time. 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 a long period.[2][4] Indeed, one species discovered in China, fossilized by fine grained volcanic ash around 165 million years ago, is hardly discernible from its modern-day descendant and belongs to the extant family of harvestman Sclerosomatidae.[11][12]

Etymology

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, United Kingdom (although not formally describing them, being a pre-Linnean work).[13]

Systematics

Main article: Harvestman phylogeny

The family Stygophalangiidae (1 species, Stygophalangium karamani) from underground waters in Macedonia is sometimes misplaced in the Phalangioidea. It is not a harvestman.

Fossil record

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

The oldest known harvestman, from the 400 million years old Devonian Rhynie chert, already has almost all the characteristics of modern species, placing the origin of harvestmen in the Silurian, or even earlier.

Interestingly, no fossils of 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 million years ago.

Naturally, most finds are from comparatively recent times. More than 20 fossil species are known from the Cenozoic, three from the Mesozoic,[12] and at least seven from the Paleozoic.[14]

Paleozoic

The 400 million years old Eophalangium sheari is known from two specimens, one a female, the other a male. The female bears an ovipositor and is about 10 millimetres (0.39 in) long, the male penis can be discerned too. It is not definitely known whether both sexes belong to the same species. They have long legs, tracheae, and no median eyes. Together with the 305 million years old Hastocularis argus, it forms the suborder Tetrophthalmi.[3][15]

Brigantibunum listoni from East Kirkton near Edinburgh in Scotland is almost 340 million years old. Its placement is rather uncertain, apart from it being a harvestman.

From about 300 million years ago (mya) there are several finds from the Coal Measures of North America and Europe.[3][4] While the two described Nemastomoides species are currently grouped as Dyspnoi, they look more like Eupnoi.

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

There are some fossils from the Permian that are possibly harvestmen, but these are not well preserved.

Described species

Mesozoic

No fossil harvestmen are known from the Triassic. They are also so far absent from the Lower Cretaceous Crato Formation of Brazil, which has yielded many other terrestrial arachnids. An unnamed long-legged harvestman was reported from the Early Cretaceous of Koonwarra, Victoria, Australia, which may be a Eupnoi.

Halitherses grimaldii from Burmese amber (c. 100 million years ago) is a long-legged Dyspnoi with large eyes, which may be related to the Ortholasmatinae (Nemastomatidae).[16]

Cenozoic

Unless otherwise noted, all species are from the Eocene.

References

  1. ↑ Adriano B. Kury (2011). "Order Opiliones Sundevall, 1833" (PDF). In Z.-Q. Zhang. Animal biodiversity: an outline of higher-level classification and survey of taxonomic richness. Zootaxa 4138. pp. 112–114.
  2. 1 2 3 4 5 6 7 8 9 Glauco Machado, Ricardo Pinto-da-Rocha & Gonzalo Giribet (2007). "What are harvestmen?". In Ricardo Pinto-da-Rocha, Glauco Machado & Gonzalo Giribet. Harvestmen: the Biology of Opiliones. Harvard University Press. pp. 1–13. ISBN 0-674-02343-9.
  3. 1 2 3 Garwood, Russell J.; Sharma, Prashant P.; Dunlop, Jason A.; Giribet, Gonzalo (2014). "A Paleozoic Stem Group to Mite Harvestmen Revealed through Integration of Phylogenetics and Development". Current Biology 24 (9): 1017–1023. doi:10.1016/j.cub.2014.03.039. Retrieved April 17, 2014.
  4. 1 2 3 Garwood, Russell J.; Dunlop, Jason A.; Giribet, Gonzalo; Sutton, Mark D. (2011). "Anatomically modern Carboniferous harvestmen demonstrate early cladogenesis and stasis in Opiliones". Nature Communications 2: 444. doi:10.1038/ncomms1458. Retrieved June 8, 2015.
  5. ↑ J. W. Shultz (1990). "Evolutionary morphology and phylogeny of Arachnida". Cladistics 6: 1–38. doi:10.1111/j.1096-0031.1990.tb00523.x.
  6. ↑ Joyce Tavolacci, ed. (2003), Insects and spiders of the world, Volume 5: Harvester ant to leaf-cutting ant, Marshall Cavendish, p. 263, ISBN 978-0-7614-7334-3
  7. ↑ Ricardo Pinto-da-Rocha and Adriano B. Kury (2003). "Third species of Guasiniidae (Opiliones, Laniatores) with comments on familial relationships" (PDF). Journal of Arachnology 31 (3): 394–399. doi:10.1636/H02-59.
  8. ↑ http://www.senckenberg.de/root/index.php?page_id=5210&year=2012&action=press&id=2480
  9. ↑ May R. Berenbaum (30 September 2009). The Earwig's Tail: a modern bestiary of multi-legged legends. Harvard University Press. p. 143. ISBN 978-0-674-05356-4.
  10. ↑ The Spider Myths Site: Daddy-Longlegs
  11. ↑ Sid Perkins (June 23, 2009). "Long-lasting daddy longlegs". Science News.
  12. 1 2 Diying Huang, Paul A. Selden & Jason A. Dunlop (2009). "Harvestmen (Arachnida: Opiliones) from the Middle Jurassic of China" (PDF). Naturwissenschaften 96 (8): 955–962. doi:10.1007/s00114-009-0556-3. PMID 19495718.
  13. ↑ Martin Lister's English Spiders, 1678. Ed. John Parker and Basil Hartley (1992). Colchester, Essex: Harley Books. pp. 26 & 30. (Translation of the Latin original, Tractatus de Araneis.)
  14. ↑ Jason A. Dunlop (2007). "Paleontology". In Ricardo Pinto-da-Rocha, Glauco Machado & Gonzalo Giribet. Harvestmen: the Biology of Opiliones. Harvard University Press. pp. 247–265. ISBN 0-674-02343-9.
  15. ↑ Fabien Tepper (April 11, 2014). "Ancient four-eyed wonder resolves daddy longleg mystery". Christian Science Monitor. Retrieved April 16, 2014.
  16. ↑ Gonzalo Giribet & Jason A. Dunlop (2005). "First identifiable Mesozoic harvestman (Opiliones: Dyspnoi) from Cretaceous Burmese amber". Proceedings of the Royal Society B 272 (1567): 1007–1013. doi:10.1098/rspb.2005.3063.

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