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''Gorgonophontes'' is an extinct
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial n ...
of
mantis shrimp Mantis shrimp, or stomatopods, are carnivorous marine crustaceans of the order Stomatopoda (). Stomatopods branched off from other members of the class Malacostraca around 340 million years ago. Mantis shrimp typically grow to around in length ...
that lived during the late Carboniferous period in what is now the United States and Belgium. It contains two named species. The type species, ''G. peleron'', was described in 1984 by
Frederick Schram Frederick Robert Schram (born August 11, 1943, in Chicago, Illinois) is an American palaeontologist and carcinologist. He received his B.S. in biology from Loyola University Chicago in 1965, and a Ph.D. on palaeozoology from the University of Ch ...
based on 100 specimens found in
Nebraska Nebraska () is a state in the Midwestern region of the United States. It is bordered by South Dakota to the north; Iowa to the east and Missouri to the southeast, both across the Missouri River; Kansas to the south; Colorado to the so ...
and
Iowa Iowa () is a U.S. state, state in the Midwestern United States, Midwestern region of the United States, bordered by the Mississippi River to the east and the Missouri River and Big Sioux River to the west. It is bordered by six states: Wiscon ...
. A second species, ''G. fraiponti'', was first named from multiple specimens found near Liège in 1922 and later reassigned to the genus. A small
crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean gro ...
with a total length of around , this animal had a subrectangular
carapace A carapace is a dorsal (upper) section of the exoskeleton or shell in a number of animal groups, including arthropods, such as crustaceans and arachnids, as well as vertebrates, such as turtles and tortoises. In turtles and tortoises, the und ...
, a
telson The telson () is the posterior-most division of the body of an arthropod. Depending on the definition, the telson is either considered to be the final segment of the arthropod body, or an additional division that is not a true segment on accou ...
that narrows into a spike at the end, and four pairs of
maxilliped An appendage (or outgrowth) is an external body part, or natural prolongation, that protrudes from an organism's body. In arthropods, an appendage refers to any of the homologous body parts that may extend from a body segment, including ant ...
s enlarged into
raptorial The term ''raptorial'' implies much the same as ''predatory'' but most often refers to modifications of an arthropod's foreleg that make it function for the grasping of prey while it is consumed, where the gripping surfaces are formed from the o ...
appendages, the frontmost pair being larger than the others. It has been suggested to be a
benthopelagic The demersal zone is the part of the sea or ocean (or deep lake) consisting of the part of the water column near to (and significantly affected by) the seabed and the benthos. The demersal zone is just above the benthic zone and forms a layer o ...
predator, using these appendages to capture prey, possibly from above, while swimming over the seabed.


Discovery and naming

The first fossils of ''Gorgonophontes'' to be discovered were found decades before the genus was named. In 1922,
Victor van Straelen Victor van Straelen (14 June 1889 – 29 February 1964) was a Belgian conservationist, palaeontologist and carcinologist. Van Straelen was born in Antwerp on 14 June 1889, and worked chiefly as a palaeontologist until his retirement in 1954. H ...
described nine fossil specimens made up mostly of
pyritized Permineralization is a process of fossilization of bones and tissues in which mineral deposits form internal casts of organisms. Carried by water, these minerals fill the spaces within organic tissue. Because of the nature of the casts, permineral ...
material on
black shale Shale is a fine-grained, clastic sedimentary rock formed from mud that is a mix of flakes of clay minerals (hydrous aluminium phyllosilicates, e.g. kaolin, Al2 Si2 O5( OH)4) and tiny fragments (silt-sized particles) of other minerals, especiall ...
, which were collected from lower Westphalian-aged deposits near Liège, Belgium. He believed they represented three new species, which he named ''
Perimecturus ''Perimecturus'' is an extinct genus of mantis shrimp that lived during the Early Carboniferous period in what is now Scotland and the United States. The first known specimens were collected near the River Esk, Dumfries and Galloway, River Esk in ...
fraiponti'', '' Palaeocaris lohesti'' and ''Anthracomysis rostrata'' respectively. These specimens were sent to the collection of the
University of Liège The University of Liège (french: Université de Liège), or ULiège, is a major public university of the French Community of Belgium based in Liège, Wallonia, Belgium. Its official language is French. As of 2020, ULiège is ranked in the 301� ...
paleontology museum, where they remain today. Although van Straelen drew figures of the fossils he assigned to each species, these figures do not adequately reflect the anatomical features seen in the actual specimens. Later, all the specimens initially studied by van Straelen would be discovered to actually represent only one species. ''Palaeocaris lohesti'' and ''Anthracomysis rostrata'' were therefore declared as
junior synonyms The Botanical and Zoological Codes of nomenclature treat the concept of synonymy differently. * In botanical nomenclature, a synonym is a scientific name that applies to a taxon that (now) goes by a different scientific name. For example, Linnae ...
of ''Perimecturus fraiponti''. In 1962, Harold Kelly Brooks reassigned this species to ''
Archaeocaris ''Archaeocaris'' (meaning "ancient shrimp") is an extinct genus of mantis shrimp that lived in North America during the Early Carboniferous period. Though it was placed as a member of the family Perimecturidae until 2008, it is currently deemed ...
'', renaming it as ''Archaeocaris fraiponti''. The genus ''Gorgonophontes'' was first named in 1984 by
Frederick Schram Frederick Robert Schram (born August 11, 1943, in Chicago, Illinois) is an American palaeontologist and carcinologist. He received his B.S. in biology from Loyola University Chicago in 1965, and a Ph.D. on palaeozoology from the University of Ch ...
, who initially established it as a
monotypic In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unispec ...
genus containing only the type species, ''G. peleron''. He reported that 100 specimens of ''Gorgonophontes'' had been found in the upper
Pennsylvanian Pennsylvanian may refer to: * A person or thing from Pennsylvania * Pennsylvanian (geology) The Pennsylvanian ( , also known as Upper Carboniferous or Late Carboniferous) is, in the International Commission on Stratigraphy, ICS geologic timesca ...
-aged black shales of
Nebraska Nebraska () is a state in the Midwestern region of the United States. It is bordered by South Dakota to the north; Iowa to the east and Missouri to the southeast, both across the Missouri River; Kansas to the south; Colorado to the so ...
and
Iowa Iowa () is a U.S. state, state in the Midwestern United States, Midwestern region of the United States, bordered by the Mississippi River to the east and the Missouri River and Big Sioux River to the west. It is bordered by six states: Wiscon ...
, occurring in the
Oread In Greek mythology, an Oread (; grc, Ὀρειάς, Oreiás, stem grc, Ὀρειάδ-, Oreiád-, label=none, la, Oreas/Oread-, from grc, ὄρος, , mountain, label=none; french: Oréade) or Orestiad (; grc, Ὀρεστιάδες, Orest ...
,
Dennis Dennis or Denis is a first or last name from the Greco-Roman name Dionysius, via one of the Christian saints named Dionysius. The name came from Dionysus, the Greek god of ecstatic states, particularly those produced by wine, which is someti ...
and Swope formations. The
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of seve ...
specimen, SDSNH 4393, was collected from the Stark Shale member of the Dennis Formation near
Papillion, Nebraska Papillion is a city in Sarpy County in the state of Nebraska, United States. Designated as the county seat, it developed as an 1870s railroad town and suburb of Omaha. The city is part of the larger five-county metro area of Omaha. Papillion's ...
. The generic name references the
Gorgons A Gorgon ( /ˈɡɔːrɡən/; plural: Gorgons, Ancient Greek: Γοργών/Γοργώ ''Gorgṓn/Gorgṓ'') is a creature in Greek mythology. Gorgons occur in the earliest examples of Greek literature. While descriptions of Gorgons vary, the te ...
from
Greek mythology A major branch of classical mythology, Greek mythology is the body of myths originally told by the ancient Greeks, and a genre of Ancient Greek folklore. These stories concern the origin and nature of the world, the lives and activities of ...
, while the specific name is derived from the Greek word ''pelor'' meaning 'monster'. In the same publication, Schram states that examination of the ''Perimecturus fraiponti'' material has found it to actually be a member of the genus ''
Tyrannophontes ''Tyrannophontes'' is an extinct genus of mantis shrimp that lived during the late Carboniferous period in what is now the Mazon Creek fossil beds of Illinois. It is the only genus in the family Tyrannophontidae. The type species, ''T. theridion' ...
'', renaming it as ''Tyrannophontes fraiponti''. However, in 2004 Lothar Schöllmann found that the species was actually more similar to ''Gorgonophontes'' than to ''Tyrannophontes'', thus renaming it again as ''Gorgonophontes fraiponti''. Later, Schram publishes a paper in 2007 reexamining Paleozoic stomatopods in which he independently reached the same conclusion that ''Gorgonophontes'' is the proper genus for this species.


Description

''Gorgonophontes'' was a
crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean gro ...
of moderate size, measuring approximately in total length. Antennae were present on the head, though these are poorly preserved, with only the scaphocerite (the large
exopod The arthropod leg is a form of jointed appendage of arthropods, usually used for walking. Many of the terms used for arthropod leg segments (called podomeres) are of Latin origin, and may be confused with terms for bones: ''coxa'' (meaning hip, plu ...
of the antenna) well-preserved in the fossils of ''G. peleron''. The
carapace A carapace is a dorsal (upper) section of the exoskeleton or shell in a number of animal groups, including arthropods, such as crustaceans and arachnids, as well as vertebrates, such as turtles and tortoises. In turtles and tortoises, the und ...
had a subrectangular shape, with short "wings" extending posteriorly on its sides. The last three
thorax The thorax or chest is a part of the anatomy of humans, mammals, and other tetrapod animals located between the neck and the abdomen. In insects, crustaceans, and the extinct trilobites, the thorax is one of the three main divisions of the ...
segments are not covered by the carapace and instead left exposed. The
maxilliped An appendage (or outgrowth) is an external body part, or natural prolongation, that protrudes from an organism's body. In arthropods, an appendage refers to any of the homologous body parts that may extend from a body segment, including ant ...
s on the second to fifth thorax segments are enlarged and
raptorial The term ''raptorial'' implies much the same as ''predatory'' but most often refers to modifications of an arthropod's foreleg that make it function for the grasping of prey while it is consumed, where the gripping surfaces are formed from the o ...
, with the pair on the second segment being most enlarged (about 1.34 times larger than the other pairs) while those on the third to fifth segments are equal in size. In ''G. peleron'', the propodi (penultimate segment) of these maxillipeds each have two rows of spines running adjacent to each other; one row is made up of closely spaced, small spines, while the other is composed of articulated large spines spaced further apart. The narrow
dactylus The dactylus is the tip region of the tentacular club of cephalopods and of the leg of some crustaceans (see arthropod leg). In cephalopods, the dactylus is narrow and often characterized by the asymmetrical placement of suckers (i.e., the ven ...
(final segment of the appendage) is believed to have rested between these two rows when retracted. Contrastingly, the maxillipedal propodi of ''G. fraiponti'' only have one row of spines, alternating between small spines and occasional large spines, as well as longitudinal keels. The segments of the
abdomen The abdomen (colloquially called the belly, tummy, midriff, tucky or stomach) is the part of the body between the thorax (chest) and pelvis, in humans and in other vertebrates. The abdomen is the front part of the abdominal segment of the tors ...
each have a slight ridge and furrow on their margins, and extend forward. A pair of small longitudinal keels are see. on the sixth
pleon The decapod (crustaceans such as a crab, lobster, shrimp or prawn) is made up of 20 body segments grouped into two main body parts: the cephalothorax and the pleon (abdomen). Each segment may possess one pair of appendages, although in various g ...
segment. Similar ridges are also seen on the
telson The telson () is the posterior-most division of the body of an arthropod. Depending on the definition, the telson is either considered to be the final segment of the arthropod body, or an additional division that is not a true segment on accou ...
, as well as a median keel on its
proximal Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position ...
half. The telson is subtriangular in shape, narrowing into a spike at the end. The protopod (basal segment) of the
uropod Uropods are posterior appendages found on a wide variety of crustaceans. They typically have functions in locomotion. Definition Uropods are often defined as the appendages of the last body segment of a crustacean. An alternative definition sugge ...
is rectangular and has two movable spines on its posterior margin, one of which is slightly curved and two to three times longer than the other. The
endopod The arthropod leg is a form of jointed appendage of arthropods, usually used for walking. Many of the terms used for arthropod leg segments (called podomeres) are of Latin origin, and may be confused with terms for bones: ''coxa'' (meaning hip, plur ...
of the uropod narrows towards the end and is only half the length of the exopod.


Classification

In his 1984 publication naming ''Gorgonophontes'',
Frederick Schram Frederick Robert Schram (born August 11, 1943, in Chicago, Illinois) is an American palaeontologist and carcinologist. He received his B.S. in biology from Loyola University Chicago in 1965, and a Ph.D. on palaeozoology from the University of Ch ...
assigned the genus to the family Tyrannophontidae, which at the time was the only established family of
Paleozoic The Paleozoic (or Palaeozoic) Era is the earliest of three geologic eras of the Phanerozoic Eon. The name ''Paleozoic'' ( ;) was coined by the British geologist Adam Sedgwick in 1838 by combining the Greek words ''palaiós'' (, "old") and ' ...
mantis shrimps other than Perimecturidae. However, in 2007 Schram revised the relations between early
stomatopod Mantis shrimp, or stomatopods, are carnivorous marine crustaceans of the order Stomatopoda (). Stomatopods branched off from other members of the class Malacostraca around 340 million years ago. Mantis shrimp typically grow to around in length ...
s and established the family Gorgonophontidae, to which ''Gorgonophontes'' was reassigned. Gorgonophontidae was erected as a
monotypic In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unispec ...
family, with ''Gorgonophontes'' as its only included genus. Later in 2009, '' Chabardella'' would be described and placed into this family as its second genus. Smith ''et al.'' (2023) conducted a
phylogenetic analysis In biology Biology is the scientific study of life. It is a natural science with a broad scope but has several unifying themes that tie it together as a single, coherent field. For instance, all organisms are made up of cells that pro ...
to determine the relations between fossil mantis shrimps, which recovered ''Gorgonophontes'' to be a
monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic ...
genus, forming a
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 clade including '' Daidal schoellmanni'' and ''Daidal acanthcercus''. Meanwhile, the family Gorgonophontidae was found to be
polyphyletic A polyphyletic group is an assemblage of organisms or other evolving elements that is of mixed evolutionary origin. The term is often applied to groups that share similar features known as homoplasies, which are explained as a result of conver ...
. The authors suggest that such a result may be due to the fragmentary nature of ''Daidal pattoni''. The results of the analysis are displayed in the
cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to ...
below:


Palaeobiology

''Gorgonophontes'' would have been a
carnivore A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose food and energy requirements derive from animal tissues (mainly muscle, fat and other ...
, based on its enlarged
raptorial The term ''raptorial'' implies much the same as ''predatory'' but most often refers to modifications of an arthropod's foreleg that make it function for the grasping of prey while it is consumed, where the gripping surfaces are formed from the o ...
appendages and the diets of other mantis shrimps. It has been theorized to be a
benthopelagic The demersal zone is the part of the sea or ocean (or deep lake) consisting of the part of the water column near to (and significantly affected by) the seabed and the benthos. The demersal zone is just above the benthic zone and forms a layer o ...
predator, swimming a little above the seabed while it hunted by grabbing its prey from above with its raptorial
maxilliped An appendage (or outgrowth) is an external body part, or natural prolongation, that protrudes from an organism's body. In arthropods, an appendage refers to any of the homologous body parts that may extend from a body segment, including ant ...
s. This is supported by the fact that its walking appendages were too short to facilitate a wide stance, and that the animal was unable to lift the front of its body, making it incapable of catching prey while standing on the bottom. Similar conditions are seen in ''
Tyrannophontes ''Tyrannophontes'' is an extinct genus of mantis shrimp that lived during the late Carboniferous period in what is now the Mazon Creek fossil beds of Illinois. It is the only genus in the family Tyrannophontidae. The type species, ''T. theridion' ...
'' and '' Tyrannosculda'', suggesting early mantis shrimps were not
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 "t ...
animals like their modern counterparts. However, the
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. Th ...
e of modern mantis shrimps are also benthopelagic predators, and it has thus been proclaimed that studying them could allow better understanding of early mantis shrimps.


References


External links

{{taxonbar, from1=Q124348104 Carboniferous arthropods of Europe Carboniferous arthropods of North America Stomatopoda Fossil taxa described in 1984 Prehistoric crustacean genera Prehistoric mantis shrimps Pennsylvanian animals of Europe Pennsylvanian animals of North America Pennsylvanian genera