Galeaspida is an extinct taxon of jawless marine and freshwater fish

Fish

Fish are a paraphyletic group of organisms that consist of all gill-bearing aquatic vertebrate animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish, as well as various extinct related groups...

. Their name is derived from a Latin word for helmet, galea, and refers to their massive bone shield on the head. Galeaspida lived in shallow, fresh water and marine environments during the Silurian

Silurian

The Silurian is a geologic period and system that extends from the end of the Ordovician Period, about 443.7 ± 1.5 Mya , to the beginning of the Devonian Period, about 416.0 ± 2.8 Mya . As with other geologic periods, the rock beds that define the period's start and end are well identified, but the...

 and Devonian

Devonian

The Devonian is a geologic period and system of the Paleozoic Era spanning from the end of the Silurian Period, about 416.0 ± 2.8 Mya , to the beginning of the Carboniferous Period, about 359.2 ± 2.5 Mya...

 times (430 to 370 million years ago) of what is now Southern China

China

Chinese civilization may refer to:* China for more general discussion of the country.* Chinese culture* Greater China, the transnational community of ethnic Chinese.* History of China* Sinosphere, the area historically affected by Chinese culture...

, Tibet

Tibet

Tibet is a plateau region in Asia, north-east of the Himalayas. It is the traditional homeland of the Tibetan people as well as some other ethnic groups such as Monpas, Qiang, and Lhobas, and is now also inhabited by considerable numbers of Han and Hui people...

 and Vietnam

Vietnam

Vietnam – sometimes spelled Viet Nam , officially the Socialist Republic of Vietnam – is the easternmost country on the Indochina Peninsula in Southeast Asia. It is bordered by China to the north, Laos to the northwest, Cambodia to the southwest, and the South China Sea –...

. Their morphology

Morphology (biology)

In biology, morphology is a branch of bioscience dealing with the study of the form and structure of organisms and their specific structural features....

 is superficially similar to that of the Heterostraci

Heterostraci

Heterostraci is an extinct class of jawless vertebrate that lived primarily in marine and estuary environments. They arose during the Ordovician, and all, save for the Psammosteids, became extinct during the late Devonian...

 more than the Osteostraci

Osteostraci

The class Osteostraci was a group of bony-armored jawless fish, termed "ostracoderms", that lived in what is now North America, Europe and Russia from the Middle Silurian to Late Devonian....

, as there is currently no evidence that the galeaspids had paired fins. However, the Galeaspida are regarded as being more closely related to the Osteostraci than Heterostraci as the morphology of the braincase is more similar to that of Osteostraci than Heterostraci.

Morphology

The defining characteristic of all galeaspids was a large opening on the dorsal

Dorsum (biology)

In anatomy, the dorsum is the upper side of animals that typically run, fly, or swim in a horizontal position, and the back side of animals that walk upright. In vertebrates the dorsum contains the backbone. The term dorsal refers to anatomical structures that are either situated toward or grow...

 surface of the head shield, which was connected to the pharynx

Pharynx

The human pharynx is the part of the throat situated immediately posterior to the mouth and nasal cavity, and anterior to the esophagus and larynx. The human pharynx is conventionally divided into three sections: the nasopharynx , the oropharynx , and the laryngopharynx...

 and gill chamber. It may have served both the olfaction

Olfaction

Olfaction is the sense of smell. This sense is mediated by specialized sensory cells of the nasal cavity of vertebrates, and, by analogy, sensory cells of the antennae of invertebrates...

 and the intake of the respiratory water similar to the nasopharyngeal duct

Duct (anatomy)

In anatomy and physiology, a duct is a circumscribed channel leading from an exocrine gland or organ.-Types of ducts:Examples include:-Duct system:...

 of hagfish

Hagfish

Hagfish, the clade Myxini , are eel-shaped slime-producing marine animals . They are the only living animals that have a skull but not a vertebral column. Along with lampreys, hagfish are jawless and are living fossils whose next nearest relatives include all vertebrates...

es. Galeaspids are also the vertebrates which have the largest number of gill

Gill

A gill is a respiratory organ found in many aquatic organisms that extracts dissolved oxygen from water, afterward excreting carbon dioxide. The gills of some species such as hermit crabs have adapted to allow respiration on land provided they are kept moist...

s, as some species of the order

Order (biology)

In scientific classification used in biology, the order is# a taxonomic rank used in the classification of organisms. Other well-known ranks are life, domain, kingdom, phylum, class, family, genus, and species, with order fitting in between class and family...

 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

Order (biology)

In scientific classification used in biology, the order is# a taxonomic rank used in the classification of organisms. Other well-known ranks are life, domain, kingdom, phylum, class, family, genus, and species, with order fitting in between class and family...

 Eugaleaspidiformes, which comprises the genera Sinogaleaspis, Meishanaspis, and Anjianspis, and the family

Family (biology)

In biological classification, family is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, class, order, genus, and species, with family fitting between order and genus. As for the other well-known ranks, there is the option of an immediately lower rank, indicated by the...

 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

Pituriaspida

The Pituriaspida are a small group of extinct armored jawless fishes with tremendous nose-like rostrums, which lived in the marine, deltaic environments of Middle Devonian Australia...

 and Osteostraci

Osteostraci

The class Osteostraci was a group of bony-armored jawless fish, termed "ostracoderms", that lived in what is now North America, Europe and Russia from the Middle Silurian to Late Devonian....

 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

Llandovery

Llandovery is a market town in Carmarthenshire, Wales, lying on the River Tywi and the A40 road.The town is served by Llandovery railway station, where there is a park and ride to Llanelli and Shrewsbury via the Heart of Wales Line...

 Epoch

Epoch (geology)

An epoch is a subdivision of the geologic timescale based on rock layering. In order, the higher subdivisions are periods, eras and eons. We are currently living in the Holocene epoch...

 of the Silurian, about 436 million years ago. During the transition from the Llandovery to the Wenlock

Wenlock

Wenlock may refer to:Places* Little Wenlock, Shropshire, England* Wenlock Edge, limestone escarpment near Much Wenlock* Much Wenlock, Shropshire, England** Wenlock ** Much Wenlock and Severn Junction railway...

, 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

Devonian

The Devonian is a geologic period and system of the Paleozoic Era spanning from the end of the Silurian Period, about 416.0 ± 2.8 Mya , to the beginning of the Carboniferous Period, about 359.2 ± 2.5 Mya...

.

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 of yet undescribed species and genus from the Fammenian epoch of the Late Devonian, found in association with the tetrapod Sinostega

Sinostega

Sinostega is an extinct genus of early tetrapod from the Late Devonian of China.The fossil was discovered in the Ningxia Hui Autonomous Region, northwest China, and consist of a fragmentary lower jawbone measuring 7 cm in length. It is the first Devonian tetrapod to be found in Asia.-References:*...

and the antiarch placoderm Remigolepis, in strata from the Northern Chinese province of Ningxia

Ningxia

Ningxia, formerly transliterated as Ningsia, is an autonomous region of the People's Republic of China. Located in Northwest China, on the Loess Plateau, the Yellow River flows through this vast area of land. The Great Wall of China runs along its northeastern boundary...

.