Agnostida is an

order Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of ...

arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and cuticle made of chiti ...

which have classically been seen as a group of highly modified

trilobites Trilobites (; meaning "three lobes") are extinct marine arthropods that form the class Trilobita. Trilobites form one of the earliest-known groups of arthropods. The first appearance of trilobites in the fossil record defines the base of the A ...

, though some recent research has doubted this placement. Regardless, they appear to be close relatives as part of the

Artiopoda The Artiopoda is a grouping of extinct arthropods that includes trilobites and their close relatives. It was erected by Hou and Bergström in 1997 to encompass a wide diversity of arthropods that would traditionally have been assigned to the Tril ...

. They are present in the

Lower Cambrian The Cambrian Period ( ; sometimes symbolized Ꞓ) was the first geological period of the Paleozoic Era, and of the Phanerozoic Eon. The Cambrian lasted 53.4 million years from the end of the preceding Ediacaran Period 538.8 million years ago ( ...

fossil record along with

from the Redlichiida,

Corynexochida Corynexochida is an order of trilobite that lived from the Lower Cambrian to the Late Devonian. Like many of the other trilobite orders, Corynexochida contains many species with widespread characteristics. The middle region of the cephalon (the ...

, and

Ptychopariida Ptychopariida is a large, heterogeneous order of trilobite containing some of the most primitive species known. The earliest species occurred in the second half of the Lower Cambrian, and the last species did not survive the Ordovician–Silurian ...

orders, and were highly diverse throughout the

Cambrian The Cambrian Period ( ; sometimes symbolized Ꞓ) was the first geological period of the Paleozoic Era, and of the Phanerozoic Eon. The Cambrian lasted 53.4 million years from the end of the preceding Ediacaran Period 538.8 million years ago ...

. Agnostidan diversity severely declined during the Cambrian-

Ordovician The Ordovician ( ) is a geologic period and system, the second of six periods of the Paleozoic Era. The Ordovician spans 41.6 million years from the end of the Cambrian Period million years ago (Mya) to the start of the Silurian Period Mya. T ...

transition, and the last agnostidans went

extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...

in the Late Ordovician.

Systematics

The Agnostida are divided into two suborders — Agnostina and

Eodiscina Eodiscina is trilobite suborder. The Eodiscina first developed near the end of the Lower Cambrian period (late Atdabanian) and became extinct at the end of the Middle Cambrian. Species are tiny to small, and have a thorax of two or three segments. ...

— which are then subdivided into a number of families. As a group, agnostids are isopygous, meaning their

pygidium The pygidium (plural pygidia) is the posterior body part or shield of crustaceans and some other arthropods, such as insects and the extinct trilobites. In groups other than insects, it contains the anus and, in females, the ovipositor. It is compo ...

is similar in size and shape to their

cephalon Cephalon, Inc. was an American biopharmaceutical company co-founded in 1987 by pharmacologist Frank Baldino, Jr., neuroscientist Michael Lewis, and organic chemist James C. Kauer—all three former scientists with the DuPont Company. Baldino s ...

. Most agnostid species were eyeless. The systematic position of the order Agnostida within the class Trilobita remains uncertain, and there has been continuing debate whether they are trilobites or a

stem group In phylogenetics, the crown group or crown assemblage is a collection of species composed of the living representatives of the collection, the most recent common ancestor of the collection, and all descendants of the most recent common ancestor. ...

. The challenge to the status has focused on Agnostina partly due to the juveniles of one genus have been found with legs differing dramatically from those of adult trilobites, suggesting they are not members of the lamellipedian

clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English ter ...

, of which trilobites are a part. Instead, the limbs of agnostids closely resemble those of stem group crustaceans, although they lack the proximal endite, which defines that group. The study suggested that they were likely the

sister taxon In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...

to the crustacean stem lineage, and, as such, part of the clade,

Crustaceomorpha Crustaceomorpha is a proposed clade of arthropods that includes crustaceans and numerous extinct groups. Synapomorphies for the clade are that the larval antenna is a feeding or locomotory organ, and there are six endopodal podomeres in post-an ...

. Other researchers have suggested, based on a

cladistic Cladistics (; ) is an approach to biological classification in which organisms are categorized in groups ("clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is typically shared derived char ...

analyses of

dorsal Dorsal (from Latin ''dorsum'' ‘back’) may refer to: * Dorsal (anatomy), an anatomical term of location referring to the back or upper side of an organism or parts of an organism * Dorsal, positioned on top of an aircraft's fuselage * Dorsal c ...

exoskeletal features, that Eodiscina and Agnostida are closely united, and the Eodiscina descended from the trilobite order

. A 2019 study of adult specimens with preserved soft tissue from the

Burgess Shale The Burgess Shale is a fossil-bearing deposit exposed in the Canadian Rockies of British Columbia, Canada. It is famous for the exceptional preservation of the soft parts of its fossils. At old (middle Cambrian), it is one of the earliest fo ...

found that agnostidans shared morphological similarities to trilobites and other related artiopodans like nektaspids, and their placement as stem-crustaceans was unsupported. The study recovered agnostidans as the sister group to trilobites within the Artiopoda.

Ecology

Scientists have long debated whether the agnostids lived a

pelagic The pelagic zone consists of the water column of the open ocean, and can be further divided into regions by depth (as illustrated on the right). The word ''pelagic'' is derived . The pelagic zone can be thought of as an imaginary cylinder or w ...

or a

benthic The benthic zone is the ecological region at the lowest level of a body of water such as an ocean, lake, or stream, including the sediment surface and some sub-surface layers. The name comes from ancient Greek, βένθος (bénthos), meaning " ...

lifestyle. Their lack of eyes, a morphology not well-suited for swimming, and their fossils found in association with other benthic trilobites suggest a benthic (bottom-dwelling) mode of life. They are likely to have lived on areas of the ocean floor which received little or no light and fed on

detritus In biology, detritus () is dead particulate organic material, as distinguished from dissolved organic material. Detritus typically includes the bodies or fragments of bodies of dead organisms, and fecal material. Detritus typically hosts comm ...

which descended from upper layers of the sea to the bottom. Their wide geographic dispersion in the

fossil record A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...

is uncharacteristic of benthic animals, suggesting a pelagic existence. The thoracic segment appears to form a hinge between the head and pygidium allowing for a bivalved

ostracod Ostracods, or ostracodes, are a class of the Crustacea (class Ostracoda), sometimes known as seed shrimp. Some 70,000 species (only 13,000 of which are extant) have been identified, grouped into several orders. They are small crustaceans, typic ...

an-type lifestyle. The orientation of the thoracic appendages appears ill-suited for benthic living. Recent work suggests that some agnostids were benthic predators, engaging in cannibalism and possibly pack-hunting behavior. They are sometimes preserved within the voids of other organisms, for instance within empty

hyolith Hyoliths are animals with small conical shells, known as fossils from the Palaeozoic era. They are at least considered as lophotrochozoan, and possibly being lophophorates, a group which includes the brachiopods, while others consider them as b ...

conchs, within

sponge Sponges, the members of the phylum Porifera (; meaning 'pore bearer'), are a basal animal clade as a sister of the diploblasts. They are multicellular organisms that have bodies full of pores and channels allowing water to circulate throu ...

s, worm tubes and under the carapaces of bivalved arthropods, presumably in order to hide from predators or strong storm currents; or maybe whilst scavenging for food. In the case of the tapering worm tubes '' Selkirkia'', trilobites are always found with their heads directed towards the opening of the tube, suggesting that they reversed in; the absence of any moulted carapaces suggests that moulting was not their primary reason for seeking shelter.

References

External links

Order Agnostida
by Sam Gon III.

by Sam Gon III.

by Larry O'Hanlon, news.discovery.com. {{Taxonbar, from=Q134109 Trilobite orders Cambrian trilobites Ordovician trilobites Fossil taxa described in 1864 Cambrian first appearances Late Ordovician extinctions Taxa named by John William Salter