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Ricinulei

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Ricinulei
Temporal range: Late Carboniferous–Recent
Cryptocellus goodnighti
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Subphylum: Chelicerata
Class: Arachnida
Order: Ricinulei
Thorell, 1876
Family: Ricinoididae
Ewing, 1929
Exant genera

For fossil genera, see text

Ricinulei is a small order of arachnids. Like most arachnids, they are predatory; eating small arthropods. They occur today in west-central Africa (Ricinoides) and the Americas (Cryptocellus and Pseudocellus) from South America to as far north as Texas, where they either inhabit leaf-litter or caves. As of 2022, 103 extant species of ricinuleids have been described worldwide, all in the single family Ricinoididae.[1] In older works they are sometimes referred to as Podogona. Due to their obscurity they do not have a proper common-name, though in academic literature they are occasionally referred to as hooded tickspiders.

In addition to the three living genera, Ricinulei has a fossil-record spanning over 300 million years, including fossils from the Late Carboniferous of Euramerica and the Cretaceous Burmese amber.

Anatomy and physiology

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The most important general account of ricinuleid anatomy remains the 1904 monograph by Hans Jacob Hansen and William Sørensen.[2] Useful further studies can be found in, e.g., the work of Pittard and Mitchell,[3] Gerald Legg[4][5] and L. van der Hammen.[6]

Ricinoides atewa nymph from Ghana

Body

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Ricinulei are typically about 5 to 10 millimetres (0.2 to 0.4 in) long. The largest Ricinulei known to ever exist was the Late Carboniferous Curculioides bohemondi with a body length of 21.77 mm (0.857 in).[7] The cuticle (or exoskeleton) of both the legs and body is remarkably thick.[8] Their most notable feature is a "hood" (or cucullus) which can be raised and lowered over the head. When lowered, it covers the mouth and the chelicerae. Living ricinuleids have no eyes, although two pairs of lateral eyes can be seen in fossils and even living species retain light-sensitive areas of cuticle in this position.

The heavy-bodied abdomen (or opisthosoma) exhibits a narrow pedicel, or waist, where it attaches to the prosoma. Curiously, there is a complex coupling mechanism between the prosoma and opisthosoma. The front margin of the opisthosoma tucks into a corresponding fold at the back of the carapace. The advantages of this unusual system are not well understood, and since the genital opening is located on the pedicel (another rather unusual feature) the animals have to 'unlock' themselves in order to mate. The abdomen is divided dorsally into a series of large plates or tergites, each of which is subdivided into a median and lateral plate.

Male Ricinoides karschii from Campo Reserve, Cameroon

Appendages

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The mouthparts, or chelicerae, are composed of two segments forming a fixed and a moveable digit. Sensory organs are also found associated with the mouthparts;[9] presumably for tasting the food. The chelicerae can be retracted and at rest they are normally hidden beneath the cucullus.

Ricinuleid pedipalps are complex appendages. They are typically used to manipulate food items, but also bear many sensory structures and are used as 'short range' sensory organs.[10] The pedipalps end in pincers that are small relative to their bodies, when compared to those of the related orders of scorpions and pseudoscorpions. Similar pincers on the pedipalps have now been found in the extinct order Trigonotarbida (see Relationships).

As in many harvestmen, the second pair of legs is longest in ricinuleids and these limbs are used to feel ahead of the animal, almost like antennae. If the pedipalps are 'short range' sensory organs, the second pair of legs are the corresponding 'long range' ones. Sensilla on the tarsi at the ends of legs I and II (which are used more frequently to sense the surroundings) differ from those of legs III and IV.[11][12] In male ricinuleids, the third pair of legs are uniquely modified to form copulatory organs. The shape of these organs is very important for taxonomy and can be used to tell males of different species apart.[13]

Internal anatomy

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An older summary of ricinuleid internal anatomy was published by Jacques Millot.[14] The midgut has been described,[15] while the excretory system consists of Malpighian tubules and a pair of coxal glands. Female ricinuleids have spermathecae,[16] presumably to store sperm. The male genitalia, sperm cells and sperm production have also been intensively studied.[17][18] Gas exchange takes place through trachea, and opens through a single pair of spiracles on the prosoma.[19] At least one Brazilian species appears to have a plastron, which may help it prevent getting wet and allow it to continue to breathe, even if inundated with water.[20]

Behavior and life history

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Male Pseudocellus pearsei from Grutas Tzabnah, Yucatán, Mexico

Ricinuleids inhabit the leaf litter of rainforest floors, as well as caves, where they search for prey with their elongate sensory second leg pair.[21] Ricinulei feed on other small invertebrates, although details of their natural prey are sparse.[22] Relatively little is known about their courtship and mating habits,[23] but males have been observed using their modified third pair of legs to transfer a spermatophore to the female. The eggs are carried under the mother's hood, until the young hatch into six-legged larva, which later molt into their eight-legged adult forms. The six-legged larva is a feature they share with Acari (see Relationships). Despite the scarce number of studies about the biology of this group, recent studies have reported nocturnal habits, as well as novel behaviors for this group, which include interactions between individuals different than mating.[24] Ricinuleids are often found in large congregations, the exact purpose of which is unknown.[25]

Fossil record

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Ricinulei are unique among arachnids in that the first one to be discovered was a fossil, described in 1837 by the noted English geologist William Buckland;[26] albeit misinterpreted as a beetle. Further fossil species were added in subsequent years by, among others, Samuel Hubbard Scudder, Reginald Innes Pocock and Alexander Petrunkevitch.

Fifteen of the twenty species of fossil ricinuleids discovered so far originate from the late Carboniferous (Pennsylvanian) coal measures of Europe and North America. They were revised in detail in 1992 by Paul Selden,[27] who placed them in a separate suborder, Palaeoricinulei.

The fossils are divided into four families: Curculioididae, Poliocheridae, Primoricinuleidae, and Sigillaricinuleidae. The poliocherids are more like modern ricinuleids in having an opisthosoma with a series of three large, divided tergites. Curculioidids, by contrast, have an opisthosoma without obvious tergites, but with a single median sulcus; a dividing line running down the middle of the back. This superficially resembles the elytra of a beetle and explains why Buckland originally misidentified the first fossil species. Five species: ?Poliochera cretacea, Primoricinuleus pugio, Hirsutisoma acutiformis, H. bruckschi, H. grimaldii and H. dentata, are known from the Cenomanian (~99 million years old) Burmese amber of Myanmar;[28][29][30][31] Curculioides bohemondi, the largest of all Ricinulei, was a member of the Curculioididae.[7] Monooculricinuleus incisus and M. semiglobosus from Burmese amber were originally described as members of Ricinulei, but they might belong to Opiliones instead.[32]

Some Carboniferous genera of Palaeoricinulei exceed modern Ricinulei in size, with bodies 24 millimetres (0.94 in) in length, and many appear to have had eyes, unlike modern representatives which are completely blind. It is likely they had a surface dwelling ecology, unlike that of modern Ricinulei.[33] The fossil genera from the Cretaceous Burmese amber are referred to the extinct order Primoricinulei, and are thought to have had a different ecology than modern species as tree-dwelling predators that crawled on bark.[31]

Genera

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As of September 2022, the World Ricinulei Catalog accepts the following genera:[34]

  • Ricinoididae Ewing, 1929 (103+ species)
  • † Curculioididae Cockerell, 1916 (12 species, Carboniferous)
  • † Hirsutisomidae Wunderlich, 2017 (4 species, Burmese amber)
  • † Poliocheridae Scudder, 1884 (5 species, Carboniferous, ?Burmese amber)
  • † Primoricinuleidae Wunderlich, 2015 (1 species, Burmese amber)
  • † Sigillaricinuleidae Wunderlich, 2022 (1 species, Burmese amber)

Relationships

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Early work

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In 1665, Robert Hooke described a large crab-like mite he observed with a microscope, he published a description of it in his book; Micrographia.[35] The first living ricinuleid described using Linnaean taxonomy was from West Africa by Félix Édouard Guérin-Méneville in 1838,[36] i.e. one year after the first fossil. This was followed by a second living example collected by Henry Walter Bates in Brazil and described by John Obadiah Westwood in 1874,[37] and a third from Sierra Leone by Tamerlan Thorell in 1892.[38] In these early studies ricinuleids were thought to be unusual harvestmen (Opiliones), and in his 1892 paper Thorell introduced the name "Ricinulei" for these animals as a suborder of the harvestman. Ricinuleids were subsequently recognized as an arachnid order in their own right in the 1904 monograph by Hansen & Soerensen. These authors recognised a group called "Arachnida micrura", comprising spiders, whip spiders, whip scorpions and ricinuleids, which they defined as having a rather narrow join between the prosoma and opisthosoma and a small 'tail end' to the opisthosoma.

Ricinuleids and mites

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Morphological studies of arachnid relationships have largely concluded that ricinuleids are most closely related to Acari (mites and ticks) though more recent phylogenomic studies refute this.[39][40] L. van der Hammen placed ricinuleids in a group called "Cryptognomae",[41] together with the anactinotrichid mites only. Peter Weygoldt and Hannes Paulus referred to ricinuleids and all mites as "Acarinomorpha".[42][43] Jeffrey Shultz used the name "Acaromorpha".[44][45] This hypothesis recognizes that both ricinuleids and mites hatch with a larval stage with only six legs, rather than the usual eight seen in arachnids. The additional pair of legs appears later during development. Some authors have also suggested that the gnathosoma, a separate part of the body bearing the mouthparts, is also a unique character for ricinuleids and mites,[46] but this feature is rather complex and difficult to interpret and other authors would restrict the presence of a gnathosoma sensu stricto to mites only.

Ricinuleids and trigonotarbids

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In 1892, Ferdinand Karsch suggested that ricinuleids were the last living descendants of the extinct arachnid order Trigonotarbida.[47] This hypothesis was widely overlooked, but was reintroduced by Jason Dunlop in 1996.[48] Characteristics shared by ricinuleids and trigonotarbids include the division of the tergites on the opisthososma into median and lateral plates and the presence of an unusual 'locking mechanism' between the two halves of the body. A further study subsequently recognised that the tip of the pedipalp in both ricinuleids and trigonotarbids ends in a similar small claw.[49] Ricinuleids as sister group of trigonotarbids was also recovered in the 2002 study by Gonzalo Giribet and colleagues.[50]

Phylogenomic studies

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Recent phylogenomic studies have recovered different relationships than those previously suggested. An analysis in early 2019 suggested the sister group of the ricinuleids may be Xiphosura, the arthropod order containing horseshoe crabs.[39] In response to this work, a more recent study placed Ricinulei and Opiliones as sister taxa.[51]

References

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  1. ^ Valdez-Mondragón, Alejandro; Juárez-Sánchez, Alma R. (2021-02-23). "A new epigean species of ricinuleid of the genus Pseudocellus (Arachnida: Ricinulei: Ricinoididae) from a tropical sub-deciduous forest in Oaxaca, Mexico". The Journal of Arachnology. 48 (3). doi:10.1636/JoA-S-20-014. ISSN 0161-8202. S2CID 232021020.
  2. ^ Hans Jacob Hansen & William Sørensen (1904). On two orders of Arachnida. Cambridge University Press. pp. 1 182.
  3. ^ Kay Pittard & Robert W. Mitchell (1972). "Comparative morphology of the life stages of Cryptocellus pelaezi (Arachnida, Ricinulei)". Graduate Studies. 1. Texas Tech University: 3–77.
  4. ^ Gerald Legg (1976). "The external morphology of a new species of ricinuleid (Arachnida) from Sierra Leone". Journal of Zoology. 59 (1): 1–58. doi:10.1111/j.1096-3642.1976.tb01007.x.
  5. ^ Gerald Legg (1976). "The external morphology of immature stages of Ricinoides karschi (Arachnida: Ricinulei)". Bulletin of the British Arachnological Society. 3: 243–248.
  6. ^ L. van der Hammen (1979). "Comparative studies in Chelicerata I. The Cryptognomae (Ricinulei, Architarbi and Anactinotrichida)". Zoologische Verhandelingen. 174 (1): 1–62.
  7. ^ a b Niall Whalen, Paul Selden. "A new, giant ricinuleid (Arachnida, Ricinulei), from the Pennsylvanian of Illinois, and the identification of a new, ontogenetically stable, diagnostic character". Journal of Paleontology, Volume 95, Issue 3, May 2021, pp. 601 - 612 DOI: https://doi.org/10.1017/jpa.2020.104
  8. ^ J. H. Kennaugh (1968). "An examination of the cuticle of three species of Ricinulei (Arachnida)". Journal of Zoology. 156 (3): 393–404. doi:10.1111/j.1469-7998.1968.tb04361.x.
  9. ^ G. Talarico, J. G. Palacios-Vargas & G. Alberti (2008). "Taste while chewing? Sensory structures in the chelicerae of Pseudocellus pearsei (Chamberlin & Ivie, 1938) (Ricinulei, Arachnida)". Revista Ibérica de Aracnología. 15: 47–53.
  10. ^ G. Talarico, J. G. Palacios-Vargas & G. Alberti (2008). "The pedipalp of Pseudocellus pearsei (Ricinulei, Arachnida) – ultrastructure of a multifunctional organ". Arthropod Structure & Development. 37 (6): 511–521. doi:10.1016/j.asd.2008.02.001. PMID 18502688.
  11. ^ Giovanni Talarico, Jose G. Palacios-Vargas, Mariano Fuentes Silva & Gerd Alberti (2005). "First ultrastructural observations on the tarsal pore organ of Pseudocellus pearsei and P. boneti (Arachnida, Ricinulei)". Journal of Arachnology. 33 (2): 604–612. doi:10.1636/04-110.1. JSTOR 4129861. S2CID 86221977.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  12. ^ Giovanni Talarico, José G. Palacios-Vargas, Mariano Fuentes Silva & Gerd Alberti (2008). "Ultrastructure of tarsal sensilla and other integument structures of two Pseudocellus species (Ricinulei, Arachnida)". Journal of Morphology. 267 (4): 441–463. doi:10.1002/jmor.10415. PMID 16425267. S2CID 25580911.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  13. ^ S. L. Tuxen (1974). "The African genus Ricinoides (Arachnida, Ricinulei)" (PDF). Journal of Arachnology. 1: 85–106.
  14. ^ Jacques Millot (1945). "L'anatomie interne des Ricinulei". Annales des Sciences Naturelles, Zoologie (in French). 7: 1–29.
  15. ^ Mario Ludwig; José G. Palacios-Vargas; Gerd Alberti (1994). "Cellular details of the midgut of Cryptocellus boneti (Arachnida: Ricinulei)". Journal of Morphology. 220 (3): 263–270. doi:10.1002/jmor.1052200305. PMID 29865385. S2CID 46930292.
  16. ^ P. M. Brignoli (1973). "On some Ricinulei of Mexico with notes on the female genital apparatus (Arachnida, Ricinulei)". Accademia Nazionale dei Lincei. 171: 153–174.
  17. ^ Gerd Alberti & José G. Palacios-Vargas (1984). "Fine structure of spermatogenesis and mature spermatozoa in Cryptocellus boneti Bolivar y Pieltain, 1941 (Arachnida, Ricinulei)". Journal of Ultrastructure Research. 87 (1): 1–12. doi:10.1016/S0022-5320(84)90111-4.
  18. ^ G. Talarico, L. F. García Hernández & P. Michalik (2008). "The male genital system of the New World Ricinulei (Arachnida): ultrastructure of spermatozoa and spermiogenesis with special emphasis on its phylogenetic implications". Arthropod Structure & Development. 37 (5): 396–409. Bibcode:2008ArtSD..37..396T. doi:10.1016/j.asd.2008.01.006. PMID 18539528.
  19. ^ Ax, Peter (2000). "Ricinulei — Acari". Multicellular Animals. pp. 130–135. doi:10.1007/978-3-662-10396-8_36. ISBN 978-3-642-08681-6.
  20. ^ Joachim Adis, Benjamin Messner & Norman Platnick (1999). "Morphological structures and vertical distribution in the soil indicate facultative plastron respiration in Cryptocellus adisi (Arachnida, Ricinulei) from Central Amazonia". Studies on Neotropical Fauna and Environment. 34 (1): 1–9. Bibcode:1999SNFE...34....1A. doi:10.1076/snfe.34.1.1.8915.
  21. ^ Joachim U. Adis, Norman I. Platnick, José W. de Morais & José M. Gomes Rodrigues (1989). "On the abundance and ecology of Ricinulei (Arachnida) from Central Amazonia, Brazil". Journal of the New York Entomological Society. 97 (2): 133–140. JSTOR 25009750.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  22. ^ J. A. L. Cooke (1967). "Observations on the biology of Ricinulei (Arachnida) with descriptions of two new species of Cryptocellus". Journal of Zoology. 151 (1): 31–42. doi:10.1111/j.1469-7998.1967.tb02864.x.
  23. ^ Gerald Legg (1977). "Sperm transfer and mating in Ricinoides hanseni (Ricinulei: Arachnida)". Journal of Zoology. 182 (1): 51–61. doi:10.1111/j.1469-7998.1977.tb04140.x.
  24. ^ García, L. F.; Torrado-León, E.; Talarico, G.; Peretti, A. V. (2015-07-01). "First Characterization of the Behavioral Repertory in a Ricinuleid: Cryptocellus narino Platnick & Paz 1979 (Arachnida, Ricinulei, Ricinoididae)". Journal of Insect Behavior. 28 (4): 447–459. doi:10.1007/s10905-015-9517-1. hdl:11336/7938. ISSN 1572-8889. S2CID 2409069.
  25. ^ Whalen, Niall; Selden, Paul (May 2021). "A new, giant ricinuleid (Arachnida, Ricinulei), from the Pennsylvanian of Illinois, and the identification of a new, ontogenetically stable, diagnostic character". Journal of Paleontology. 95 (3): 601–612. doi:10.1017/jpa.2020.104. ISSN 0022-3360. S2CID 233301949.
  26. ^ Buckland, William (1837). Geology and mineralogy with reference to natural theology. The Bridgewater treatises on the power, wisdom and goodness of God as manifested in the creation. Vol. Treatise IV (2nd ed.). London, UK: William Pickering.
  27. ^ Selden, P.A. (1992). "Revision of the fossil ricinuleids". Transactions of the Royal Society of Edinburgh. Earth Sciences. 83 (4): 595–634. doi:10.1017/s0263593300003333. S2CID 85945454.
  28. ^ Wunderlich, Jörg (2012). "Description of the first fossil Ricinulei in amber from Burma (Myanmar), the first report of this arachnid order from the Mesozoic and from Asia, with notes on the related extinct order Trigonotarbida". In Wunderlich, Jörg (ed.). Beiträge zur Araneologie. Vol. 7: Fifteen papers on extant and fossil spiders (Araneae). pp. 233–244.
  29. ^ Wunderlich, Jörg (2015). "New and rare fossil Arachnida in Cretaceous Burmese amber (Amblypygi, Ricinulei and Uropygi: Thelephonida)". In Wunderlich, Jörg (ed.). Beiträge zur Araneologie. Vol. 9: Mesozoic spiders and other fossil arachnids. pp. 409–436.
  30. ^ Wunderlich, Jörg (2017). "New extinct taxa of the arachnid order Ricinulei, based on new fossils preserved in mid Cretaceous Burmese amber". In Wunderlich, Jörg (ed.). Beiträge zur Araneologie. Vol. 10. pp. 48–71.
  31. ^ a b Botero-Trujillo, Ricardo; Davis, Steven R.; Michalik, Peter; Prendini, Lorenzo (22 September 2022). "Hirsutisoma grimaldii sp. nov., a ca. 99 million-year-old ricinuleid (Primoricinulei, Hirsutisomidae) from Cretaceous Burmese amber with a corticolous, scansorial lifestyle". Palaeoentomology. 5 (5). doi:10.11646/palaeoentomology.5.5.11. ISSN 2624-2834.
  32. ^ Selden, Paul A.; Ren, Dong (2017). "A review of Burmese amber arachnids". Journal of Arachnology. 45 (3): 324–343. doi:10.1636/JoA-S-17-029. S2CID 90983791.
  33. ^ Whalen, Niall; Selden, Paul (May 2021). "A new, giant ricinuleid (Arachnida, Ricinulei), from the Pennsylvanian of Illinois, and the identification of a new, ontogenetically stable, diagnostic character". Journal of Paleontology. 95 (3): 601–612. doi:10.1017/jpa.2020.104. ISSN 0022-3360.
  34. ^ "World Ricinulei Catalog". World Ricinulei Catalog. Natural History Museum Bern. 2022. Retrieved 24 September 2022.
  35. ^ Hooke, Robert (1665). "Of the crab-like insect". Micrographia, or Some physiological descriptions of minute bodies made by magnifying glasses with observations and inquiries thereupon. London: James Allestry and John Martyn in the Royal Society. pp. 207–208.
  36. ^ Félix Édouard Guérin-Méneville (1838). "Note sur l'Acanthodon et sur le Cryptostemme, nouveaux genres d'Arachnides". Revue Zoologique par la Société Cuvierienne (in French). 1: 10–12.
  37. ^ John Obadiah Westwood (1874). "Class Arachnida". Thesaurus Entomologicus Oxoniensis. Oxford: Clarendon Press. pp. 200–202.
  38. ^ Tamerlan Thorell (1892). "On an apparently new arachnid belonging to the family Cryptostemmoidae, Westw". Kungliga Svenska Ventenskaps-akademiens Handlingar. 17: 1–18.
  39. ^ a b Ballesteros, Jesús A.; Sharma, Prashant P. (2019). "A Critical Appraisal of the Placement of Xiphosura (Chelicerata) with Account of Known Sources of Phylogenetic Error". Systematic Biology. 68 (6): 896–917. doi:10.1093/sysbio/syz011. PMID 30917194.
  40. ^ Lozano-Fernandez, L.; et al. (2019). "Increasing species sampling in chelicerate genomic-scale datasets provides support for monophyly of Acari and Arachnida". Nature Communications. 10 (2295): 459–462. Bibcode:2019NatCo..10.2295L. doi:10.1038/s41467-019-10244-7. PMC 6534568. PMID 6534568.
  41. ^ L. van der Hammen (1977). "A new classification of Chelicerata". Zoologische Mededelingen. 51 (20): 307–319.
  42. ^ Peter Weygoldt & Hannes Paulus (1979). "Untersuchungen zur Morphologie, Taxonomie und Phylogenie der Chelicerata. I. Morphologische Untersuchungen". Zeitschrift für zoologische Systematik und Evolutionsforschung (in German). 17 (3): 85–116. doi:10.1111/j.1439-0469.1979.tb00694.x.
  43. ^ Peter Weygoldt & Hannes Paulus (1979). "Untersuchungen zur Morphologie, Taxonomie und Phylogenie der Chelicerata. II. Cladogramme und die Entfaltung der Chelicerata". Zeitschrift für zoologische Systematik und Evolutionsforschung (in German). 17 (3): 177–200. doi:10.1111/j.1439-0469.1979.tb00699.x.
  44. ^ Jeffrey W. Shultz (1990). "Evolutionary morphology and phylogeny of Arachnida". Cladistics. 6 (1): 1–38. doi:10.1111/j.1096-0031.1990.tb00523.x. PMID 34933471. S2CID 85410687.
  45. ^ Jeffrey W. Shultz (2007). "A phylogenetic analysis of the arachnid orders based on morphological characters". Zoological Journal of the Linnean Society. 150 (2): 221–265. doi:10.1111/j.1096-3642.2007.00284.x.
  46. ^ E. E. Lindquist (1984). "Current theories on the evolution of major groups of Acari and on their relationships with other groups of Arachnida with consequent implications for their classification". In D. A. Griffiths; C. E. Bowman (eds.). Acarology VI, Volume 1. Chichester: Ellis Horwood Ltd. pp. 28–62. ISBN 978-0-85312-603-4.
  47. ^ Ferdinand Karsch (1892). "Ueber Cryptostemma Guèr. als einziger recenter Ausläufer der fossilen Arachnoideen-Ordnung Meridogastra Thor". Berliner Entomologische Zeitschrift (in German). 37 (1): 25–32. doi:10.1002/mmnd.18920370108.
  48. ^ Jason A. Dunlop (1996). "Evidence for a sister group relationship between Ricinulei and Trigonotarbida" (PDF). Bulletin of the British Arachnological Society. 10 (6): 193–204. Archived from the original (PDF) on 2011-06-13. Retrieved 2010-11-11.
  49. ^ Jason A. Dunlop, Carsten Kamenz and Giovanni Talarico (2009). "A fossil trigonotarbid arachnid with a ricinuleid-like pedipalpal claw". Zoomorphology. 128 (4): 305–313. doi:10.1007/s00435-009-0090-z. S2CID 6769463.
  50. ^ Gonzalo Giribet, Gregory D. Edgecombe, Ward C. Wheeler & Courtney Babbitt (2002). "Phylogeny and systematic position of Opiliones: a combined analysis of chelicerate relationships using morphological and molecular data" (PDF). Cladistics. 18 (1): 5–70. doi:10.1111/j.1096-0031.2002.tb00140.x. PMID 14552352. S2CID 16833833.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  51. ^ Lozano-Fernandez, L.; et al. (2019). "Increasing species sampling in chelicerate genomic-scale datasets provides support for monophyly of Acari and Arachnida". Nature Communications. 10 (2295): 459–462. Bibcode:2019NatCo..10.2295L. doi:10.1038/s41467-019-10244-7. PMC 6534568. PMID 6534568.

Further reading

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