Triarthrus Becki
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''Triarthrus'' is a genus of
Upper Ordovician The Late Ordovician is the third and final epoch of the Ordovician period, lasting million years and spanning from around 458.2 to 443.1 million years ago. The rocks associated with this epoch are referred to as the Upper Ordovician Series. At ...
ptychopariid
trilobite Trilobites (; meaning "three-lobed entities") are extinction, extinct marine arthropods that form the class (biology), class Trilobita. One of the earliest groups of arthropods to appear in the fossil record, trilobites were among the most succ ...
found in
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,
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,
Kentucky Kentucky (, ), officially the Commonwealth of Kentucky, is a landlocked U.S. state, state in the Southeastern United States, Southeastern region of the United States. It borders Illinois, Indiana, and Ohio to the north, West Virginia to the ...
, and
Indiana Indiana ( ) is a U.S. state, state in the Midwestern United States, Midwestern region of the United States. It borders Lake Michigan to the northwest, Michigan to the north and northeast, Ohio to the east, the Ohio River and Kentucky to the s ...
, eastern and northern
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, China and
Scandinavia Scandinavia is a subregion#Europe, subregion of northern Europe, with strong historical, cultural, and linguistic ties between its constituent peoples. ''Scandinavia'' most commonly refers to Denmark, Norway, and Sweden. It can sometimes also ...
. It is the last of the Olenid trilobites, a group which flourished in the
Cambrian The Cambrian ( ) is the first geological period of the Paleozoic Era, and the Phanerozoic Eon. The Cambrian lasted 51.95 million years from the end of the preceding Ediacaran period 538.8 Ma (million years ago) to the beginning of the Ordov ...
period. The specimens of ''T. eatoni'' that are found in the
Beecher's Trilobite Bed Beecher's Trilobite Bed is a Konservat-Lagerstätte of Late Ordovician (Caradoc) age located within the Frankfort Shale in Cleveland's Glen, Oneida County, New York, USA.Rome Rome (Italian language, Italian and , ) is the capital city and most populated (municipality) of Italy. It is also the administrative centre of the Lazio Regions of Italy, region and of the Metropolitan City of Rome. A special named with 2, ...
,
New York New York most commonly refers to: * New York (state), a state in the northeastern United States * New York City, the most populous city in the United States, located in the state of New York New York may also refer to: Places United Kingdom * ...
area are exquisitely preserved showing soft body parts in
iron pyrite The mineral pyrite ( ), or iron pyrite, also known as fool's gold, is an iron sulfide with the chemical formula Fe S2 (iron (II) disulfide). Pyrite is the most abundant sulfide mineral. Pyrite's metallic luster and pale brass-yellow hue ...
. Pyrite preservation has given scientists a rare opportunity to examine the
gills A gill () is a respiratory organ that many aquatic organisms use to extract dissolved oxygen from water and to excrete carbon dioxide. The gills of some species, such as hermit crabs, have adapted to allow respiration on land provided they are ...
, walking legs, antennae, digestive systems, and eggs of trilobites, which are rarely preserved. ''Triarthrus'' is therefore commonly used in science texts to illustrate trilobite anatomy and physiology.


Distribution

* ''T. beckii'' Upper Caradoc and Ashgill, Snake Hill Formation, Cohoes, New York State; and Kentucky. * ''T. billingsi'' Ashgill?, Quebec * ''T. canadensis'' is known from the Upper Ordovician of Canada (Katian, lower Member of the Whitby Formation, Craigleith vicinity, Georgian Bay area; middle Member of the Whitby Formation, Whitby, Rogue River and Pickering, all Lake Simcoe area, Ontario) ''cited in'' * ''T. eatoni'' Upper Caradoc-Ashgill, N.Y., is known from the Upper Ordovician of Canada (Ashgill, lower Member of the Whitby Formation, Craigleith vicinity, Georgian Bay area, Lake Simcoe area, Ontario; and Quebec) and the United States (New York) * ''T. glaber'' Ashgill, Quebec * ''T. huguesensis'' Ashgill, Quebec * ''T. latissimus'' Sweden * ''T. linnarssoni'' Norway and Sweden * ''T. rougensis'' Ashgill, Ontario * ''T. sichuansis'' China * ''T. spinosus'' is known from the Upper Ordovician of Canada (Katian, middle Member of the Whitby Formation, Rogue River and Pickering, both Lake Simcoe area, Ontario and Quebec), and the United States (New York). * ''T. novoaustralis'' is known from Upper Ordovician of Australia (middle Katian, Malachis Hill Formation)


Taxonomy

''T. beckii'' and ''T. eatoni'' have long been considered closely related and possibly synonymous. Recent comparative analysis showed that there is no sharp distinction between the two, but that they seem to represent opposing morphotypes. ''T. beckii'' dominates earlier in the distribution and in shallow water, while ''T. eatoni'' does so later and in deeper waters. It is probable that the transgression of the sea level aided the eventual disappearance of the ''T. beckii'' morph. The occurrence of the supposed pluriform species continued for more than two million years.


Species previously assigned to ''Triarthrus''

* ''T. angelini'' = '' Bienvillia angelini'' * ''T. belli'' = '' Parabolinella'' sp. * ''T. caenigenus'' = '' Porterfieldia caenigena'' * ''T. convergens'' = ''Porterfieldia convergens'' * ''T. fisheri'' = ''Porterfieldia fisheri'' * ''T. goldwyerensis'' = ''Porterfieldia goldwyerensis'' * ''T. humilis'' = ''Porterfieldia humilis'' * ''T. jachalensis'' = ''Porterfieldia jachalensis'' * ''T. pacificus'' = ''Porterfieldia pacifica'' * ''T. papilosus'' = ''Porterfieldia papilosa'' * ''T. parapunctatus'' = ''Porterfieldia parapunctata'' * ''T. parchaensis'' = ''Bienvillia parchaensis'' * ''T. punctatus'' = ''Porterfieldia punctata'' * ''T. rectifrons'' = ''Bienvillia rectifrons'' * ''T. reedi'' = ''Porterfieldia caenigena'' * ''T. shinetonensis'' = ''Bienvillia shinetonensis'' * ''T. sinensis'' = ''Porterfieldia sinensis'' * ''T. tetragonalis'' = ''Bienvillia tetragonalis'' * ''T. thor'' = ''Porterfieldia thor'' * ''T. variscorum'' = '' Parabolina frequens''


Description


Exoskeleton

''Triarthrus'' is an average size trilobite (up to about ) and its moderately convex body is about twice as long as wide (excluding spines). Like in all Olenidae, the headshield (or
cephalon Cephalon, Inc. was an American biopharmaceutical company co-founded in 1987 by pharmacologist Frank Baldino Jr., Frank Baldino, Jr., neuroscientist Michael Lewis, and organic chemist James C. Kauer—all three former scientists with the DuPont ...
) of ''Triarthrus'' has
opisthoparian Trilobites (; meaning "three-lobed entities") are extinct marine arthropods that form the class Trilobita. One of the earliest groups of arthropods to appear in the fossil record, trilobites were among the most successful of all early animals, ...
sutures, and the right and left free cheeks that they define are yoked. The cephalon in ''Triarthrus'' is semicircular. The central raised area (or
glabella The glabella, in humans, is the area of skin between the eyebrows and above the nose. The term also refers to the underlying bone that is slightly depressed, and joins the two brow ridges. It is a cephalometric landmark that is just superior ...
) is approximately quadrate, and considerably wider than the posterior margin of the fixed cheeks (or fixigenae). The front of the glabella is close to the anterior border (or the preglabellar field is short), but the border furrow is absent in front of the glabella. The facial suture crosses in front of the glabella on the top of an inflated rim. The posterior end of the glabella consists of the occipital ring that is defined by a furrow crossing over the midline. The occipital ring carries a small node at its centre point, and may be adorned with a backward directed spine as long as the glabella. In front of the occipital furrow two pairs of lateral furrows emerge from the axial furrow almost perpendicularly and curve backwards as they cut toward the midline, but without reaching it. Anteriorly, one or two further furrows are isolated shallow slits or depressions. The fixigenae are very narrow (less than ¼ of the width of the glabella at the back of the eye, and less than ½ at the occipital ring). The librigenae are narrow, and with or without spines. The border is prominent elsewhere, and the eyes of small to medium size, sitting at the end of small palpebral lobes next to the frontal half of the glabella. The thorax consists of 13 to 16 segments, with the axis wider than pleural regions, obliquely truncated or rounded pleural tips, and the fulcrum placed very close to the axis. The tailpiece (or
pygidium The pygidium (: 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 compos ...
) is small, with 3 to 5 axial rings, distinctly segmented pleural fields, and an entire, evenly rounded margin.


Uncalcified parts

The foremost pair of extremities in trilobites are the
antennas In radio-frequency engineering, an antenna (American English) or aerial (British English) is an electronic device that converts an alternating electric current into radio waves (transmitting), or radio waves into an electric current (receivi ...
. In ''Triarthrus eatoni'' the antennas are probably flexible. From their attachment adjacent to the
hypostome In zoology, the hypostome can refer to structures in distinct animal groups: *Hypostome (trilobite), the ventral mouthpart plate in trilobites *Hypostome (tick) The hypostome (also called the maxilla, radula, or Labium (insect), labium) is a Os ...
they stretch inward and forward, almost touching each other at the point where, from a
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 The fus ...
viewpoint, they appear from under the cephalic margin, then bending outward and then forward again in a slight S-curve. They are about twice as long as the headshield (or
cephalon Cephalon, Inc. was an American biopharmaceutical company co-founded in 1987 by pharmacologist Frank Baldino Jr., Frank Baldino, Jr., neuroscientist Michael Lewis, and organic chemist James C. Kauer—all three former scientists with the DuPont ...
), each consisting of 40 to 50 segments that are shorter than wide. As the preservation of soft tissue in ''T. eatoni'' is excellent, it is probable that it lacked cerci, the most backward pair of extremities known from '' Olenoides serratus''.


Life cycle

Due to the existence of an excellently preserved deposit in the Frankfurt Shale near Rome, New York State, that suggests an entire population of ''Triarthrus eatoni'' was killed and quickly buried, it has been possible to make a plausible reconstruction of its life history. After hatching from roughly spherical eggs averaging ~0.2mm in diameter, the small protaspid and early meraspid stages supposedly lived between the
plankton Plankton are the diverse collection of organisms that drift in Hydrosphere, water (or atmosphere, air) but are unable to actively propel themselves against ocean current, currents (or wind). The individual organisms constituting plankton are ca ...
in the
water column The (oceanic) water column is a concept used in oceanography to describe the physical (temperature, salinity, light penetration) and chemical ( pH, dissolved oxygen, nutrient salts) characteristics of seawater at different depths for a defined ...
. This is derived from the fact that only
exuvia In biology, exuviae are the remains of an exoskeleton and related structures that are left after ecdysozoans (including insects, crustaceans and arachnids) have molted. The exuviae of an animal can be important to biologists as they can often b ...
e of the early stages were found, making clear that they haven't died in the disaster that killed their elders. In this phase the larvae probably dispersed over large areas. After a few months, at about 2mm long ''T. eatoni'' started living at the
sea floor The seabed (also known as the seafloor, sea floor, ocean floor, and ocean bottom) is the bottom of the ocean. All floors of the ocean are known as seabeds. The structure of the seabed of the global ocean is governed by plate tectonics. Most of ...
. Here, it must have fed by filtering suspended food particles, probably comparable to the trunk limb filter-feeding
Cephalocarida The Cephalocarida, from Ancient Greek κεφαλή (''kephalḗ''), meaning "head", and καρίς (''karís''), meaning "shrimp", are a class in the subphylum Crustacea comprising only 12 species. Both the nauplii and the adults are benthic. ...
and
Branchiopoda Branchiopoda, from Ancient Greek βράγχια (''bránkhia''), meaning "gill", and πούς (''poús''), meaning "foot", is a class (biology), class of crustaceans. It comprises Anostraca, fairy shrimp, clam shrimp, Diplostraca (or Cladocera), ...
. The exoskeletons fall into five size categories, suggesting that ''T. eatoni'' had a distinct breeding season, likely annually, as is the case in almost all
extant Extant or Least-concern species, least concern is the opposite of the word extinct. It may refer to: * Extant hereditary titles * Extant literature, surviving literature, such as ''Beowulf'', the oldest extant manuscript written in English * Exta ...
crustacea Crustaceans (from Latin meaning: "those with shells" or "crusted ones") are invertebrate animals that constitute one group of arthropods that are traditionally a part of the subphylum Crustacea (), a large, diverse group of mainly aquatic arthrop ...
ns (although some
amphipod Amphipoda () is an order of malacostracan crustaceans with no carapace and generally with laterally compressed bodies. Amphipods () range in size from and are mostly detritivores or scavengers. There are more than 10,700 amphipod species cur ...
s have two breeding periods per year). It could live through at least four breeding seasons, and at that age reach approximately 4 cm in length.


References


Sources


''T. eatoni'' and ''T. rougensis'' photos.


Further reading

*Thomas E. Whiteley, Gerald J. Kloc, and Carlton E. Brett ''Trilobites of New York''. Cornell University Press, 2002. 456 pages. {{Taxonbar, from=Q606700 Olenina Ptychopariida genera Ordovician trilobites of North America Paleozoic life of Ontario Paleozoic life of the Northwest Territories Paleozoic life of Quebec Ordovician trilobites of Europe