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In 2024, a study discovered that living members of
Opiliones have retained lateral eyes, as well as a second pair of vestigial median eyes. A reanalysis of harvestman phylogeny based on 78 genes and an updated understanding of morphology revealed that Tetrophthalmi are derived members of the living
241:(commonly known as harvestmen or daddy-longlegs) that had both median and lateral eyes. First described in 2014, it is known from two extinct species. Phylogenetic analysis suggests that this eye arrangement is the ancestral condition for harvestmen, placing Tetrophthalmi and
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The most recent phylogenetic analyses of
Opiliones phylogeny have recovered Tetrophthalmi as derived members within the suborder Eupnoi, which has resulted in older ages of Opiliones diversification than previously estimated. The restoration of
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specimen found a suppressed gene that, if active, would generate a second pair of eyes at the lateral location, providing independent evidence that four eyes is the ancestral condition. Garwood
324:(a part of the reproductive system), and external male genitalia – which distinguish them from all other harvestmen. Thus, the two species were classified into a new suborder, Tetrophthalmi.
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in Greek mythology. According to their analysis, the specimen died, and was rapidly buried approximately 305 million years ago. Before it decayed, its body was surrounded by the mineral
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and the new suborder
Tetrophthalmi help explain how arachnid eyes developed. The classification also helps clarify when side-eyed and fore-head eyed harvestmen split into distinct
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in the middle of head and one set on its side on the stink glands. Performing comparative analysis on 158 morphological features, Garwood's team determined
447:, the oldest known Opiliones fossil, to membership in Eupnoi is consistent with the original description of this species as a true daddy-longlegs.
467:
Gainett, Guilherme; Klementz, Benjamin C.; Blaszczyk, Pola; Setton, Emily V.W.; Murayama, Gabriel; Willemart, Rodrigo H.; Gavish-Regev, Efrat;
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312:. The two species were not, however, as closely related to the 270 other species of harvestmen studied. The team determined
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also argue that a
Carboniferous harvestmen diversification is more consistent with changes observed in other terrestrial
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375:. The Cyphophthalmi stem then diversified into Cyphophthalmi proper and Tetrophthalmi, while the Phalangida split into
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273:(CT scans) to investigate the fossil. In 2014, a team led by Garwood identified the specimen as a novel species,
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548:"A Paleozoic Stem Group to Mite Harvestmen Revealed through Integration of Phylogenetics and Development"
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deposit in
Eastern France. The fossil went unstudied for more than 30 years before paleontologist
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473:"Vestigial organs alter fossil placements in an ancient group of terrestrial chelicerates"
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as an early branching opilionid in the harvestman tree, along with the
Cyphophthalmi.
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Garwood, Russell J.; Sharma, Prashant P.; Dunlop, Jason A.; Giribet, Gonzalo (2014).
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s
Tetrophthalmi was recovered in a polytomy with the remaining oplionids, however.
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620:"Three-dimensional reconstruction and the phylogeny of extinct chelicerate orders"
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395:. The opiliones' own divergence is dated to 414 million years ago, and
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and the "Palpatores". Finally, the
Palpatores diversified into
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showing dorsal, lateral, ventral, and anterolateral views
515:"Ancient four-eyed wonder resolves daddy longleg mystery"
491:"Ancient four-eyed wonder resolves daddy longleg mystery"
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In the early 1980s, a tiny fossil was extracted from the
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320:shared three features – two pairs of eyes, an open
399:are estimated to have originated during the late
371:Opiliones split into the "Phalangida" and stem
618:Garwood, Russell J.; Dunlop, Jason A. (2014).
589:"Ancient Daddy Longlegs Had Extra Set of Eyes"
298:had two sets of eyes - one set mounted on the
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184:Garwood, Sharma, Dunlop & Giribet, 2014
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16:Suborder of harvestmen/daddy longlegs
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427:Another recent phylogeny recovered
346:Phylogenetic analysis of Opiliones
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587:Stefan Sirucek (April 11, 2014).
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513:Fabien Tepper (April 11, 2014).
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263:Montceau-les-Mines Lagerstätte
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435:, the other genus in Garwood
789:Prehistoric animal suborders
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249:position within Opiliones.
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294:The CT scan revealed that
565:10.1016/j.cub.2014.03.039
519:Christian Science Monitor
495:Christian Science Monitor
271:X-ray computed tomography
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119:Scientific classification
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257:Classification history
30:Devonian–Carboniferous
751:Paleobiology Database
471:(February 23, 2024).
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681:at Wikimedia Commons
352:Harvestman phylogeny
109:A reconstruction of
794:Paleozoic arachnids
784:Arthropod suborders
593:National Geographic
309:Eophalangium sheari
599:on August 31, 2014
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328:Hastocularis argus
276:Hastocularis argus
111:Hastocularis argus
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691:Taxon identifiers
677:Media related to
637:10.7717/peerj.641
445:Eophangium sheari
413:Phalangium opilio
365:phylogenetic tree
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27:Temporal range:
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24:Tetrophthalmi
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601:. Retrieved
597:the original
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723:Wikispecies
356:In Garwood
779:Harvestmen
773:Categories
451:References
422:arthropods
405:Ordivician
377:Laniatores
350:See also:
289:concretion
224:2 species
173:Suborder:
142:Arthropoda
714:Q16919670
603:April 17,
524:April 16,
403:to early
397:arachnids
338:Phylogeny
322:gonostome
318:E. sheari
300:ocularium
239:Opiliones
218:Diversity
166:Opiliones
154:Arachnida
125:Kingdom:
708:Wikidata
656:25405073
630:: e641.
574:24726154
401:Cambrian
389:Devonian
314:H. argus
304:H. argus
296:H. argus
285:siderite
235:suborder
137:Phylum:
130:Animalia
647:4232842
437:et al.'
385:Dyspnoi
190:Genera
161:Order:
149:Class:
756:294819
654:
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572:
418:et al.
381:Eupnoi
367:, the
358:et al.
332:clades
743:7NGCV
624:PeerJ
369:basal
361:'
269:used
247:basal
245:in a
652:PMID
605:2014
570:PMID
526:2014
383:and
316:and
35:Preęž’
738:CoL
642:PMC
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