Tetrophthalmi

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Tetrophthalmi
Temporal range: Devonian–Carboniferous
Reconstruction of Hastocularis argus.jpg
A reconstruction of Hastocularis argus showing dorsal, lateral, ventral, and anterolateral views
Scientific classification
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Suborder:
Tetrophthalmi

Garwood, Sharma, Dunlop & Giribet, 2014
Genera
Diversity
2 species

Tetrophthalmi is an extinct suborder of Opiliones (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 Cyphophthalmi in a basal position within Opiliones.

Contents

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 suborder Eupnoi (true daddy-longlegs). [1]

Classification history

In the early 1980s, a tiny fossil was extracted from the Montceau-les-Mines Lagerstätte deposit in Eastern France. The fossil went unstudied for more than 30 years before paleontologist Russell Garwood used X-ray computed tomography (CT scans) to investigate the fossil. In 2014, a team led by Garwood identified the specimen as a novel species, Hastocularis argus , named for the 100-eyed giant Argus Panoptes in Greek mythology. [2] 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 siderite, which formed a protective concretion. The specimen subsequently decayed, leaving a void showing a detailed outline of the creature. [3]

The CT scan revealed that H. argus had two sets of eyes - one set mounted on the ocularium 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 H. argus was closely related to Eophalangium sheari . [3] The two species were not, however, as closely related to the 270 other species of harvestmen studied. The team determined H. argus and E. sheari shared three features – two pairs of eyes, an open gonostome (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. [4]

Hastocularis argus 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 clades. [3]

Phylogeny

Phylogenetic analysis of Opiliones Phylogenetic Analyses of Opiliones 2014-A.png
Phylogenetic analysis of Opiliones

In Garwood et al.'s phylogenetic tree, the basal Opiliones split into the "Phalangida" and stem Cyphophthalmi. The Cyphophthalmi stem then diversified into Cyphophthalmi proper and Tetrophthalmi, while the Phalangida split into Laniatores and the "Palpatores". Finally, the Palpatores diversified into Eupnoi and Dyspnoi. The analysis moves the divergence of the extant suborders from the Devonian Period to the Carboniferous. The opiliones' own divergence is dated to 414 million years ago, and arachnids are estimated to have originated during the late Cambrian to early Ordivician. [4]

Genetic analysis performed on a modern Phalangium opilio 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. [5] [4] Garwood et al. also argue that a Carboniferous harvestmen diversification is more consistent with changes observed in other terrestrial arthropods, which have been linked to high oxygen levels during that period. [4]

Another recent phylogeny recovered Hastocularis as an early branching opilionid in the harvestman tree, along with the Cyphophthalmi. [6] Eophalangium, the other genus in Garwood et al.'s Tetrophthalmi was recovered in a polytomy with the remaining oplionids, however.

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. [1] The restoration of Eophangium sheari, the oldest known Opiliones fossil, to membership in Eupnoi is consistent with the original description of this species as a true daddy-longlegs.

Related Research Articles

<span class="mw-page-title-main">Chelicerata</span> Subphylum of arthropods

The subphylum Chelicerata constitutes one of the major subdivisions of the phylum Arthropoda. Chelicerates include the sea spiders, horseshoe crabs, and arachnids, as well as a number of extinct lineages, such as the eurypterids and chasmataspidids.

<span class="mw-page-title-main">Arachnid</span> Class of arthropods

Arachnida is a class of joint-legged arthropods, in the subphylum Chelicerata. Arachnida includes, among others, spiders, scorpions, ticks, mites, pseudoscorpions, harvestmen, camel spiders, whip spiders and vinegaroons.

<span class="mw-page-title-main">Opiliones</span> Order of arachnids (harvestmen/daddy longlegs)

The Opiliones are an order of arachnids colloquially known as harvestmen, harvesters, harvest spiders, or daddy longlegs. As of April 2017, 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.

<i>Plesiosiro</i> Extinct genus of arachnids

Plesiosiro is an extinct arachnid genus known exclusively from nine specimens from the Upper Carboniferous of Coseley, Staffordshire, United Kingdom. The genus is monotypic, represented only by the species Plesiosiro madeleyi described by Reginald Innes Pocock in his important 1911 monograph on British Carboniferous arachnids. It is the only known member of the order Haptopoda.

<span class="mw-page-title-main">Eupnoi</span> Suborder of harvestmen/daddy longlegs

The Eupnoi are a suborder of harvestmen, with more than 200 genera, and about 1,700 described species.

<span class="mw-page-title-main">Dyspnoi</span> Suborder of harvestmen/daddy longlegs

Dyspnoi is a suborder of harvestmen, currently comprising 43 extant genera and 356 extant species, although more species are expected to be described in the future. The eight families are currently grouped into three superfamilies: the Acropsopilionoidea, Ischyropsalidioidea, and Troguloidea.

<span class="mw-page-title-main">Cyphophthalmi</span> Suborder of harvestmen/daddy longlegs

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.

<i>Phalangium opilio</i> Species of harvestman/daddy longlegs

Phalangium opilio is a species of harvestman belonging to the family Phalangiidae.

<span class="mw-page-title-main">Caddidae</span> Family of harvestmen/daddy longlegs

Caddoidea superfamily of harvestmen arachnids with a single family Caddidae, which now only contains 2 extant species. The family previously contained many more taxa under a previous wider concept, but the familial definition was narrowed after restudy.

<span class="mw-page-title-main">Troglosironidae</span> Family of harvestmen/daddy longlegs

Troglosironidae is a family of harvestmen with seventeen described species in a single genus, Troglosiro, which is found on the island of New Caledonia, in the Pacific Ocean.

<span class="mw-page-title-main">Ogoveidae</span> Family of harvestmen/daddy longlegs

Ogoveidae is a family of harvestmen with three described species in one genus, Ogovea, which is found in equatorial West Africa.

An ozophore is an elevated cone present in the harvestman suborder Cyphophthalmi. It carries the openings, called ozopores, of the defensive glands that are present in many harvestmen.

<span class="mw-page-title-main">Ozopore</span> Opening of a defensive gland present in some arthropods

An ozopore is the opening of a defensive gland present in some arthropods, notably in millipedes of the order Polydesmida and in harvestmen, the eight-legged arachnids also known as "daddy long-legs". The glands themselves are known as ozadenes, also called "scent glands", "repugnatorial glands", "odoriferous glands" or "stink glands" by various authors. The name is derived from Ancient Greek ozo "smell" and Latin porus "pore, small opening".

Nemastomoides is an extinct genus of harvestmen known from the Carboniferous fossil record. The genus is the only member of the family Nemastomoididae and contains three described species. Nemastomoides elaveris was found in the Coal Measures of Commentry in northern France, together with Eotrogulus fayoli.

<span class="mw-page-title-main">Opiliones anatomy</span>

Opiliones are an order of arachnids and share many common characteristics with other arachnids. However, several differences separate harvestmen from other arachnid orders such as spiders. The bodies of opiliones are divided into two tagmata : the abdomen (opisthosoma) and the cephalothorax (prosoma). Unlike spiders, the juncture between the abdomen and cephalothorax is often poorly defined. Harvestmen have chelicerae, pedipalps and four pairs of legs. Most harvestmen have two eyes, although there are eyeless species.

<span class="mw-page-title-main">Harvestman phylogeny</span> Order of arachnids

Harvestmen (Opiliones) are an order of arachnids often confused with spiders, though the two orders are not closely related. Research on harvestman phylogeny is in a state of flux. While some families are clearly monophyletic, that is share a common ancestor, others are not, and the relationships between families are often not well understood.

Gonzalo Giribet is a Spanish-American invertebrate zoologist and Alexander Agassiz Professor of zoology working on systematics and biogeography at the Museum of Comparative Zoology in Harvard University. He is a past president of the International Society for Invertebrate Morphology, of the Willi Hennig Society, and vice-president of the Sociedad Española de Malacología.

<span class="mw-page-title-main">Uraraneida</span> Order of arachnids

Uraraneida is an extinct order of Paleozoic arachnids related to modern spiders. Two genera of fossils have been definitively placed in this order: Attercopus from the Devonian of United States and Permarachne from the Permian of Russia. Like spiders, they are known to have produced silk, but lack the characteristic spinnerets of modern spiders, and retain elongate telsons.

<i>Forsteropsalis photophaga</i> Species of long-legged harvestman in the family Neopilionidae

Forsteropsalis photophaga, also known as the glow-worm hunter, is a species of long-legged harvestman in the family Neopilionidae. This species is endemic to New Zealand, found in North Island caves in the vicinity of Waitomo. The name "photophaga" comes from their habit of feeding on the luminescent larvae, pupae, and adults of the New Zealand glow-worm Arachnocampa luminosa.

Prashant P. Sharma is an Indian-American invertebrate biologist and a professor of Integrative Biology at the University of Wisconsin-Madison.

References

  1. 1 2 Gainett, Guilherme; Klementz, Benjamin C.; Blaszczyk, Pola; Setton, Emily V.W.; Murayama, Gabriel; Willemart, Rodrigo H.; Gavish-Regev, Efrat; Sharma, Prashant (February 23, 2024). "Vestigial organs alter fossil placements in an ancient group of terrestrial chelicerates". Current Biology.
  2. "Ancient four-eyed wonder resolves daddy longleg mystery". Christian Science Monitor. 11 April 2014.
  3. 1 2 3 Fabien Tepper (April 11, 2014). "Ancient four-eyed wonder resolves daddy longleg mystery". Christian Science Monitor. Retrieved April 16, 2014.
  4. 1 2 3 4 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 . PMID   24726154.
  5. Stefan Sirucek (April 11, 2014). "Ancient Daddy Longlegs Had Extra Set of Eyes". National Geographic. Archived from the original on August 31, 2014. Retrieved April 17, 2014.
  6. Garwood, Russell J.; Dunlop, Jason A. (2014). "Three-dimensional reconstruction and the phylogeny of extinct chelicerate orders". PeerJ. 2: e641. doi: 10.7717/peerj.641 . PMC   4232842 . PMID   25405073.