Galeaspida

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Galeaspida
Temporal range: Llandovery [1] Early Devonian, 438–400  Ma
Galeaspida 1.JPG
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Subphylum: Vertebrata
Class: Galeaspida
Liu, 1965
Orders

Eugaleaspidiformes Liu, 1965
Huananaspidiformes Janvier, 1975
Polybranchiaspidiformes Janvier, 1996

Contents

Galeaspida (from Latin, 'Helmet shields') is an extinct taxon of jawless marine and freshwater fish. The name is derived from galea, the Latin word for helmet, and refers to their massive bone shield on the head. Galeaspida lived in shallow, fresh water and marine environments during the Silurian and Devonian times (430 to 370 million years ago) in what is now Southern China, Tibet and Vietnam. Superficially, their morphology appears more similar to that of Heterostraci than Osteostraci, there being currently no evidence that the galeaspids had paired fins.[ citation needed ] A galeaspid Tujiaaspis vividus from the Silurian period of China was described in 2022 as having a precursor condition to the form of paired fins seen in Osteostraci and gnathostomes. [2] Earlier than this, Galeaspida were already in fact regarded as being more closely related to Osteostraci, based on the closer similarity of the morphology of the braincase.[ citation needed ]

Morphology

Reconstruction of Hayangaspis (left), Dayongaspis (middle), and Changxingaspis (right) Basal galeaspid reconstructions yang.png
Reconstruction of Hayangaspis (left), Dayongaspis (middle), and Changxingaspis (right)
Their mouth and gill openings are situated on the ventral surface of the head (top right). In the most primitive forms, such as the Silurian genus Hanyangaspis (top), the median dorsal inhalent opening is broad and situated anteriorly. In other galeaspids, it is more posterior in position and can be oval, rounded, heart-shaped or slit-shaped. In some Devonian galeaspids, such as the hunanaspidiforms Lungmenshanaspis (middle) and Sanchaspis (bottom right), the headshield is produced laterally and anteriorly into slender processes. The eugaleaspidiforms, such as Eugaleaspis (bottom left) have a horseshoe-shaped headshield and a slit-shaped median dorsal opening, which imitates the aspect of the headshield of osteostracans. Galeaspida.gif
Their mouth and gill openings are situated on the ventral surface of the head (top right). In the most primitive forms, such as the Silurian genus Hanyangaspis (top), the median dorsal inhalent opening is broad and situated anteriorly. In other galeaspids, it is more posterior in position and can be oval, rounded, heart-shaped or slit-shaped. In some Devonian galeaspids, such as the hunanaspidiforms Lungmenshanaspis (middle) and Sanchaspis (bottom right), the headshield is produced laterally and anteriorly into slender processes. The eugaleaspidiforms, such as Eugaleaspis (bottom left) have a horseshoe-shaped headshield and a slit-shaped median dorsal opening, which imitates the aspect of the headshield of osteostracans.
Headshield of Nochelaspis NochelaspisMaeandrine-PaleozoologicalMuseumOfChina-May23-08.jpg
Headshield of Nochelaspis
Headshield of Laxaspis LaxaspisQujingensis-PaleozoologicalMuseumOfChina-May23-08.jpg
Headshield of Laxaspis

The defining characteristic of all galeaspids was a large opening on the dorsal surface of the head shield, which was connected to the pharynx and gill chamber, and a scalloped pattern of the sensory-lines.

The opening appears to have served both the olfaction and the intake of the respiratory water similar to the nasopharyngeal duct of hagfishes. [3] Galeaspids are also the vertebrates which have the largest number of gills, as some species of the order Polybranchiaspidida (literally "many gills shields") had up to 45 gill openings. The body is covered with minute scales arranged in oblique rows and there is no other fin besides the caudal fin. The mouth and gill openings are situated on the ventral side of the head, which is flat or flattened and suggests that they were bottom-dwellers.

Taxonomy

There are around 76 + described species of galeaspids in at least 53 genera.

If the families Hanyangaspidae and Xiushuiaspidae can be ignored as basal galeaspids, the rest of Galeaspida can be sorted into two main groups: the first being the order Eugaleaspidiformes, which comprises the genera Sinogaleaspis , Meishanaspis , and Anjianspis , and the family Eugaleaspididae, and the second being the Supraorder Polybranchiaspidida, which comprises the order Polybranchiaspidiformes, which is the sister taxon of the family Zhaotongaspididae and the order Huananaspidiformes, and the family Geraspididae, which is the sister taxon of Polybranchiaspidiformes + Zhaotongaspididae + Huananaspidiformes.

Some experts demote Galeaspida to the rank of subclass, and unite it with Pituriaspida and Osteostraci to form the class Monorhina.

Fossil record

The oldest known galeaspids, such as those of the genera Hanyangaspis and Dayongaspis , first appear near the start of the Telychian age, of the latter half of the Llandovery epoch of the Silurian, about 436 million years ago. During the transition from the Llandovery to the Wenlock, the Eugaleaspids underwent a diversification event. By the time the Wenlock epoch transitioned into the Ludlow Epoch, all of the eugaleaspids, save for the Eugaleaspidae, were extinct. The Eugaleaspidae lived from the Wenlock, and were fairly long-lived, especially the genus Eugaleaspis . The last of the Eugaleaspididae disappeared by the end of the Pragian Epoch of the Lower Devonian.

The first genus of Geraspididae, the eponymous Geraspis , appears during the middle of the Telychian. The other genera of Polybranchiaspidida appear in the fossil record a little after the beginning of the Lochkovian Epoch, at the start of the Devonian. The vast majority of the supraorder's genera either date from the Pragian epoch, or have their ranges end there. By the time the Emsian epoch starts, only a few genera, such as Duyunolepis and Wumengshanaspis , survive, with most others already extinct. The last galeaspid is an as yet undescribed species and genus from the Fammenian epoch of the Late Devonian, found in association with the tetrapod Sinostega and the antiarch placoderm Remigolepis , in strata from the Northern Chinese province of Ningxia.

Taxa

Restoration of Rhegmaspis, Dongfangaspis, and Nanningaspis Ecological restoration of galeapids.png
Restoration of Rhegmaspis , Dongfangaspis , and Nanningaspis

See also

Related Research Articles

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The Silurian is a geologic period and system spanning 24.6 million years from the end of the Ordovician Period, at 443.8 million years ago (Mya), to the beginning of the Devonian Period, 419.2 Mya. The Silurian is the shortest period of the Paleozoic Era. As with other geologic periods, the rock beds that define the period's start and end are well identified, but the exact dates are uncertain by a few million years. The base of the Silurian is set at a series of major Ordovician–Silurian extinction events when up to 60% of marine genera were wiped out.

<span class="mw-page-title-main">Agnatha</span> Infraphylum of jawless fish

Agnatha is an infraphylum of jawless fish in the phylum Chordata, subphylum Vertebrata, consisting of both living (cyclostomes) and extinct species. Among recent animals, cyclostomes are sister to all vertebrates with jaws, known as gnathostomes.

<span class="mw-page-title-main">Gnathostomata</span> Infraphylum of vertebrates

Gnathostomata are the jawed vertebrates. Gnathostome diversity comprises roughly 60,000 species, which accounts for 99% of all living vertebrates, including humans. In addition to opposing jaws, living gnathostomes have true teeth, paired appendages, the elastomeric protein of elastin, and a horizontal semicircular canal of the inner ear, along with physiological and cellular anatomical characters such as the myelin sheaths of neurons, and an adaptive immune system that has the discrete lymphoid organs of spleen and thymus, and uses V(D)J recombination to create antigen recognition sites, rather than using genetic recombination in the variable lymphocyte receptor gene.

<span class="mw-page-title-main">Sarcopterygii</span> Class of fishes

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<span class="mw-page-title-main">Cephalaspidomorphi</span> Extinct clade of jawless fishes

Cephalaspidomorphs are a group of jawless fishes named for Cephalaspis of the osteostracans. Most biologists regard this taxon as extinct, but the name is sometimes used in the classification of lampreys, because lampreys were once thought to be related to cephalaspids. If lampreys are included, they would extend the known range of the group from the Silurian and Devonian periods to the present day. They are the closest relatives of jawed fishes, who emerged from within them and they would survive if the jawed fish are included.

<span class="mw-page-title-main">Acanthodii</span> Class of fishes (fossil)

Acanthodii or acanthodians is an extinct class of gnathostomes. They are currently considered to represent a paraphyletic grade of various fish lineages basal to extant Chondrichthyes, which includes living sharks, rays, and chimaeras. Acanthodians possess a mosaic of features shared with both osteichthyans and chondrichthyans. In general body shape, they were similar to modern sharks, but their epidermis was covered with tiny rhomboid platelets like the scales of holosteians.

<span class="mw-page-title-main">Placoderm</span> Class of fishes (fossil)

Placoderms are vertebrate animals of the class Placodermi, an extinct group of prehistoric fish known from Paleozoic fossils during the Silurian and the Devonian periods. While their endoskeletons are mainly cartilaginous, their head and thorax were covered by articulated armoured plates, and the rest of the body was scaled or naked depending on the species.

<span class="mw-page-title-main">Heterostraci</span> Extinct subclass of jawless fishes

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<span class="mw-page-title-main">Anaspida</span> Group of extinct jawless vertebrates

Anaspida is an extinct group of jawless fish that existed from the early Silurian period to the late Devonian period. They were classically regarded as the ancestors of lampreys, but it is denied in recent phylogenetic analysis, although some analysis show these group would be at least related. Anaspids were small marine fish that lacked a heavy bony shield and paired fins, but were distinctively hypocercal.

<i>Minicrania</i> Extinct genus of fishes

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<i>Shuyu</i> Extinct genus of jawless fishes

Shuyu is an extinct genus of early jawless vertebrate from the Early or Middle Silurian period. It is the basalmost known eugaleaspidiform galeaspid and it lived in what is now northwestern Zhejiang Province, Southeast China. It is known from more than 20 headshields and at least 20 of them include three-dimensionally preserved neurocrania. The specimens of Shuyu were collected from the lower part of Maoshan Formation, located in Changxing District. Shuyu zhejianensis was first assigned by Pan, 1986 to a species of Sinogaleaspis. The genus was first named by Zhikun Gai, Philip C. J. Donoghue, Min Zhu, Philippe Janvier and Marco Stampanoni in 2011 and the type species is Shuyu zhejianensis.

The Xikeng Formation is located in Xiushui County, Jiangxi Province, and contains alternating beds of purplish red, grayish green and yellow green sandy and muddy rocks. The Xikeng Formation is dated to the Late Silurian - Early Devonian period.

<span class="mw-page-title-main">Evolution of fish</span> Origin and diversification of fish through geologic time

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<i>Xiangshuiosteus</i> Extinct genus of fishes

Xiangshuiosteus wui is an extinct monospecific genus of brachythoracid arthrodire placoderm from the Late Emsian stage of the Early Devonian epoch, discovered in Wuding County of Yunnan province, China. It has recently been reassessed as a dunkleosteid.

<i>Shimenolepis</i>

Shimenolepis granifera is an extinct yunnanolepid placoderm from the Xiaoxi Formation, Li County, Hunan, China. Its age is discussed, while originally considered as late Llandovery, it is later considered to belong to the Ludlow Epoch instead. It was the first described Silurian placoderm, and was the earliest known placoderm until Xiushanosteus was described, known from distinctively ordered plates.

Eugaleaspidiformes is an extinct order of jawless marine and freshwater fish, which lived in East Asia from the Telychian to the Lower Devonian period. The order was first named by Lui in 1965.

This list of fossil fish research presented in 2022 is a list of new taxa of jawless vertebrates, placoderms, acanthodians, fossil cartilaginous fishes, bony fishes, and other fishes that were described during the year, as well as other significant discoveries and events related to paleoichthyology that occurred in 2022.

References

  1. Sansom, Robert S.; Randle, Emma; Donoghue, Philip C. J. (February 7, 2015). "Discriminating signal from noise in the fossil record of early vertebrates reveals cryptic evolutionary history". Proceedings of the Royal Society B . 282 (1800): 20142245. doi:10.1098/rspb.2014.2245. PMC   4298210 . PMID   25520359.
  2. Gai, Zhikun; Li, Qiang; Ferrón, Humberto G.; Keating, Joseph N.; Wang, Junqing; Donoghue, Philip C. J.; Zhu, Min (2022-09-29). "Galeaspid anatomy and the origin of vertebrate paired appendages". Nature. 609 (7929): 959–963. doi:10.1038/s41586-022-04897-6. hdl: 1983/39f23cba-4a2b-4d8f-92fe-dbfc89869c26 . ISSN   0028-0836.
  3. Fossil jawless fish from China foreshadows early jawed vertebrate anatomy The galeaspids are characterized by a large median dorsal opening in the anterior part of the headshield that serves as both a common nostril and the main water intake device.