Phacops Plate
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''Phacops'' is a genus of
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 ...
s in the order
Phacopida Phacopida ("lens-face") is an order of trilobites that lived from the Late Cambrian to the Late Devonian. It is made up of a morphologically diverse assemblage of taxa in three related suborders. Characteristics Phacopida had 8 to 19 thoraci ...
, family
Phacopidae Phacopidae is a family (biology), family of Phacopida, phacopid trilobites that ranges from the Lower Ordovician to the Upper Devonian, with representatives in all Supercontinents, paleocontinents. Description As in all Phacopina, the eyes (if ...
, that lived in Europe, northwestern Africa, North and South America and China from the Late Ordovician until the very end of the
Devonian The Devonian ( ) is a period (geology), geologic period and system (stratigraphy), system of the Paleozoic era (geology), era during the Phanerozoic eon (geology), eon, spanning 60.3 million years from the end of the preceding Silurian per ...
, with a broader time range described from the
Late 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 ...
.''Phacops''
at
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It was a rounded animal, with a globose head and large eyes, and probably fed on
detritus In biology, detritus ( or ) is organic matter made up of the decomposition, decomposing remains of organisms and plants, and also of feces. Detritus usually hosts communities of microorganisms that colonize and decomposition, decompose (Reminera ...
. ''Phacops'' is often found rolled up ("
volvation Volvation (from Latin ''volvere'' "roll", and the suffix ''-(a)tion''; sometimes called enrolment or conglobation), is a defensive behavior in certain animals, in which the animal rolls its own body into a ball, presenting only the hardest parts ...
"), a biological defense mechanism that is widespread among smaller trilobites but further perfected in this genus.


Description

Like in all sighted
Phacopina The Phacopina comprise a suborder of the trilobite order Phacopida. Species belonging to the Phacopina lived from the Lower Ordovician (Tremadocian) through the end of the Upper Devonian (Famennian).Moore, R.C. (ed.). Treatise on Invertebrate Pal ...
, the eyes of ''Phacops'' are compounded of very large, separately set lenses without a common cornea (so called schizochroal eyes), and like almost all other Phacopina, the articulate mid-length part of the body (or
thorax The thorax (: thoraces or thoraxes) or chest is a part of the anatomy of 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 di ...
) in ''Phacops'' has 11 segments. 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 ...
) of 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 ...
) is moderately to strongly inflated near to its front, more or less flattened on the top, falling vertically to or slightly overhanging the border furrow. Up to three lateral furrows may be discernable on the glabella behind the utterly dominating frontal lobe. From the back there is a very distinct occipital ring, and in front of that a distinct preoccipital ring which is weakly divided into a strongly convex central lobe and weakly convex lateral lobes. The large to medium size eyes have a crescent shaped outline, and are elevated high above the cheeks. The steep visual surface is kidney shaped. The back corners of the cephalon (or genal angles) are acutely to bluntly rounded, but a genal spine is lacking in adults. In the ventral surface of the seam (or doublure) is in the frontal half of the cephalon a continuous furrow, delineated by ridges, and with notches laterally. This so-called vindicular furrow serves to lock the rim of the tailshield to the headshield when the trilobite is enrolled. The axial rings of the thorax do not have convex lateral axial nodes on its outer surface. The tailshield (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 well segmented. The pygidial axis has 9 to 11 rings, and the pleural areas to the sides have 5 to 8 pairs of recognizable ribs. Furrows between the ribs are deep, those that divide each rib in frontal and rear bands are very shallow, and the frontal bands are widest. The surface of the
exoskeleton An exoskeleton () . is a skeleton that is on the exterior of an animal in the form of hardened integument, which both supports the body's shape and protects the internal organs, in contrast to an internal endoskeleton (e.g. human skeleton, that ...
is covered in
tubercle In anatomy, a tubercle (literally 'small tuber', Latin for 'lump') is any round nodule, small eminence, or warty outgrowth found on external or internal organs of a plant or an animal. In plants A tubercle is generally a wart-like projectio ...
s.


Camouflage

There are specimens known of ''Phacops rana'' with many irregular black spots. Because similar spots in a specimen of '' Greenops boothi'' from the same site are arranged in rows, it may be assumed that they are original and not caused by the fossilisation process. The spots are irregular and have spurs branching outwardly, similar to the melanophores in many extant animals. In one specimen, the black spots are much larger than in another one. It is quite conceivable that changing the size of the melanophores enabled ''Phacops rana'' to camouflage itself in different environments.


Taxonomy

The concept of many fossil taxa has been tightened over time, including ''Phacops''. As a result, ''Boeckops'', ''Chotecops'', ''Paciphacops'', ''Prokops'' and ''Viaphacops'' have been erected as subgenera of ''Phacops'', and are now widely regarded as genera in their own right. Most recent, it was considered that some North-American and North-African species on the one hand and European species on the other hand differ sufficiently from each other to be assigned to separate genera. As the
type species In International_Code_of_Zoological_Nomenclature, zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the spe ...
is the European ''P. latifrons'', the North-American species are now called '' Eldredgeops''. However, the previous assigned species, like '' Phacops rana'', are still widely used among fossil collectors. ''Eldredgeops'' has a raised ridge along the ventral margin of the cephalon, the glabella is more inflated, the lateral parts of the preoccipital ring are not round but rectangular, the palpebral area and palpebral lobe are larger than in ''P. latifrons'', and there is no fold right behind the posterior vertical row of lenses, nor an isolated raised area just below the lenses. Not all of these characters may differentiate between ''Eldredgeops'' and other ''Phacops'' species however. During the
Eifelian The Eifelian is the first of two faunal stages in the Middle Devonian Epoch. It lasted from 393.3 ± 1.2 million years ago to 387.7 ± 0.8 million years ago. It was preceded by the Emsian Stage and followed by the Givetian Stage. North American ...
in the present-day Belgian
Ardennes The Ardennes ( ; ; ; ; ), also known as the Ardennes Forest or Forest of Ardennes, is a region of extensive forests, rough terrain, rolling hills and ridges primarily in Belgium and Luxembourg, extending into Germany and France. Geological ...
, several ''Phacops'' species developed from each other, the oldest being ''P. imitator'', followed by ''P. fragosus'', then ''P. latifrons'' and finally ''P. sartenaeri''. These species show a decrease in the number of lenses, which is a more widespread and recurring trend in many Phacopinae. Fossils of ''Phacops salteri'' have been found in the trilobite-rich
Late Emsian The Devonian ( ) is a geologic period and system of the Paleozoic era during the Phanerozoic eon, spanning 60.3 million years from the end of the preceding Silurian period at million years ago ( Ma), to the beginning of the succeeding Ca ...
to Early Givetian
Floresta Formation The Floresta Formation (, Df) is a geological formation of the Altiplano Cundiboyacense in the Eastern Ranges of the Colombian Andes. The sequence of siltstones, shales, coquinas and sandstone beds dates to the Devonian period; Late Emsian, Eif ...
of the
Altiplano Cundiboyacense The Altiplano Cundiboyacense () is a high plateau located in the Eastern Cordillera of the Colombian Andes covering parts of the departments of Cundinamarca and Boyacá. (Do not confuse with The Altiplano or the Altiplano Nariñense, both fur ...
, Colombia.Floresta Fauna
at
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Species

''Phacops'' currently contains the following species: * ''P. accipitrinus'' (Phillips, 1841) * ''P. algericus'' Alberti, 1983 * ''P. breviceps'' Barrande, 1846 * ''P. chlupaci'' Alberti, 1983 * ''P. circumspectans'' Paeckelmann, 1913 * ''P. degener'' Barrande, 1852 * '' P. fecundus'' * ''P. fragosus'' Struve, 1970 * ''P. granulatus'' (Munster, 1840) synonyms ''Calymene granulata'', ''P. posidoniae'' * ''P. hoseri'' Hawle & Corda, 1847 * ''P. iowensis'' Delo, 1935 * ''P. imitator'' Struve, 1970 * ''P. kockeli'' Alberti, 1968 * ''P. latifrons'' (Bronn, 1825) * ''P. maurulus'' * ''P. modestus'' Barrande * ''P. ouarouroutensis'' Crônier, 2018 * ''P. platilegnotor'' * ''P. salteri'' Kozlowski, 1923 * ''P. sartenaeri'' Struve, 1985 * ''P. sobolevi'' Kielan, 1954 * ''P. sternbergi'' * ''P. turco'' Richter & Richter * ''P. wedekindi'' Richter & Richter, 1926 * ''P. zinkeni'' F.A. Roemer, 1843


Species previously assigned to ''Phacops''

A number of species previously assigned to the genus ''Phacops'' have since been transferred to other genera: * ''P. acuticeps'' = '' Acuticryphops acuticeps'' * ''P. arcticus'' = '' Acernaspis arctica'' * ''P. asper'' = '' Ananaspis aspera'' * ''P. birdsongensis'' = ''
Paciphacops birdsongensis ''Paciphacops'' is a genus of trilobites within order Phacopida, suborder Phacopina. This genus is found primarily in the United States and Australia and is easily mistaken for the genera ''Phacops'' and ''Kainops'', which are also popular among ...
'' * ''P. boecki'' = '' Boeckops boecki'' * ''P. braziliensis'' = '' Phacopidina braziliensis'' * ''P. bronni'' = '' Reedops bronni'' * ''P. bulliceps'' = '' Eophacops bulliceps'' * ''P. caecus'' = '' Trimerocephalus caecus'' * ''P. caffer'' = ''
Metacryphaeus caffer ''Metacryphaeus'' is an extinct genus of trilobites from the family Calmoniidae. ''Metacryphaeus'' fossils have been found in Bolivia, Brazil, South Africa and Uruguay. The genus was named in 1907.F. R. C. Reed. 1907. A new species of ''Lichas ...
'' * ''P. cambelli'' = '' Paciphacops cambelli'' * ''P. claviger'' = ''
Paciphacops claviger ''Paciphacops'' is a genus of trilobites within order Phacopida, suborder Phacopina. This genus is found primarily in the United States and Australia and is easily mistaken for the genera ''Phacops'' and ''Kainops'', which are also popular among ...
'' * ''P. caudatus'' = '' Dalmanites caudatus'' * ''P. constrictus'' = '' Acastoides constrictus'' * ''P. coronatus'' = '' Heliocephalus coronatus'' * ''P. cristatus bombifrons'' = '' Viaphacops bombifrons'' * ''P. cristatus cristatus'' = '' Viaphacops cristatus'' * ''P. crossleii'' = '' Paciphacops crossleii'' * ''P. cryptophthalmoides'' = '' Trimerocephalus cryptophthalmoides'' * ''P. cryptophthalmus'' = '' Cryphops cryptophthalmus'' * ''P. cultifrons'' = '' Eophacops trapeziceps'' * ''P. dagincourti'' = '' Bouleia dagincourti'' * ''P. deshayesi'' = '' Zeliszkella deshayesi'' * ''P. downingiae'' var. ''α. vulgaris'' = '' Acaste downingiae'' * ''P. downingiae'' var. ''β. macrops'' = '' Acastocephala macrops'' * ''P. downingiae'' var. ''γ. inflatus'' = '' Acaste inflata'' * ''P. ensae'' = '' Weyerites ensae'' * ''P. fecundus'' = '' Ananaspis fecunda'' * ''P. fecundus minor'' = '' Lochkovella minor'' * ''P. ferdinandi'' = '' Chotecops ferdinandi'' * ''P. glaber'' = '' Eophacops glaber'' * ''P. glockeri'' = '' Phacopidella glockeri'' * ''P. grimbergi'' = '' Lochkovella grimbergi'' * ''P. handwerki'' = '' Eophacops handwerki'' * ''P. hanusi'' = '' Lochkovella hanusi'' * ''P. hoeninghausi'' = '' Prokops hoeninghausi'' * ''P. hudsonicus'' = '' Paciphacops hudsonicus'' * ''P. incisus'' = '' Nephranops incisus'' * ''P. khatangensis'' = '' Acernaspis khatangensis'' * ''P. kayseri'' = '' Eocryphops kayseri'' * ''P. lacunosus'' = '' Trimerocephalus lacunosus'' * ''P. latigenalis'' = ''
Paciphacops latigenalis ''Paciphacops'' is a genus of trilobites within order Phacopida, suborder Phacopina. This genus is found primarily in the United States and Australia and is easily mistaken for the genera ''Phacops'' and ''Kainops'', which are also popular among ...
'' * ''P. latilimbatus'' = '' Cryphops latilimbatus'' * ''P. lentiginosus'' = '' Trimerocephalus lentiginosus'' * ''P. limbatus'' = '' Dianops limbatus'' * ''P. logani'' = ''
Paciphacops logani ''Paciphacops'' is a genus of trilobites within order Phacopida, suborder Phacopina. This genus is found primarily in the United States and Australia and is easily mistaken for the genera ''Phacops'' and ''Kainops'', which are also popular among ...
'' * ''P. lopatini'' = '' Monorakos lopatini'' * ''P. mancus'' = '' Eophacops mancus'' * ''P. marklandensis'' = '' Acernaspis marklandensis'' * ''P. mastophthalmus'' = '' Trimerocephalus mastophthalmus'' * ''P. metacernaspis'' = generic assignment uncertain * ''P. michelini'' = '' Pseudocryphaeus michelini'' * ''P. microps'' = ''
Kainops microps ''Kainops'' is a genus of trilobites from the family Phacopidae, order Phacopida. It can be distinguished from '' Paciphacops'' by the greater number of facets to the eye (6–8 per row, compared to 3–4 in ''Paciphacops''). The form of the furr ...
'' * ''P. milleri'' = '' Eldredgeops milleri'' * ''P. miser'' = '' Lochkovella miser'' * ''P. musheni'' = '' Eophacops musheni'' * ''P. nudus'' = generic assignment uncertain * ''P. ocellus'' = '' Hadrorachus ocellus'' * ''P. opitzi'' = '' Chotecops opitzi'' * ''P. orestes'' = '' Acernaspis orestes'' * ''P. orientalis'' = '' Ananaspis orientalis'' * ''P. parabolus'' = '' Phillipsinella parabola'' * ''P. phillipsi'' = '' Kloucekia phillipsi'' * ''P. primaevus'' = '' Acernaspis primaeva'' * ''P. prokopi'' = '' Prokops prokopi'' * ''P. pulcellus'' = '' Acernaspis pulcella'' * ''P. quadrilineatus'' = '' Acernaspis quadrilineata'' * ''P. rana'' = '' Eldredgeops rana'' * ''P. raymondi'' = ''
Kainops raymondi ''Kainops'' is a genus of trilobites from the family Phacopidae, order Phacopida. It can be distinguished from '' Paciphacops'' by the greater number of facets to the eye (6–8 per row, compared to 3–4 in ''Paciphacops''). The form of the furr ...
'' * ''P. schlosseri'' = '' Struveops schlosseri'' * ''P. serratus'' = '' Paciphacops serratus'' * ''P. schlotheimi'' = '' Geesops schlotheimi'' * ''P. socialis'' = '' Dalmanitina socialis'' * ''P. speculator'' = '' Austerops speculator'' * ''P. sphaericeps'' = '' Dereimsia sphaericeps'' * ''P. steinachensis'' = '' Trimerocephalus steinachensis'' * ''P. stellifer'' = '' Greenops (Neometacanthus) stellifer'' * ''P. stokesii'' = generic assignment uncertain * ''P. straitonensis'' = '' Podowrinella straitonensis'' * ''P. trapeziceps'' = '' Eophacops trapeziceps'' * ''P. veles'' = ''
Kainops veles ''Kainops'' is a genus of trilobites from the family Phacopidae, order Phacopida. It can be distinguished from '' Paciphacops'' by the greater number of facets to the eye (6–8 per row, compared to 3–4 in ''Paciphacops''). The form of the furr ...
'' * ''P. vodorezovi'' = '' Trimerocephalus vodorezovi'' * ''P. volborthi'' = '' Denckmannites volborthi'' * ''P. wocklumeriae'' = '' Weyerites wocklumeriae''


References


External links

* {{Taxonbar, from=Q44129 Phacopidae Ordovician trilobites of Europe Silurian trilobites of Europe Devonian trilobites of Europe Ordovician United Kingdom Trilobites of North America Silurian United States Devonian United States Fossils of Georgia (U.S. state) Trilobites of South America Devonian Bolivia Devonian Brazil Devonian Colombia Fossils of Colombia Devonian Venezuela Ordovician first appearances Late Devonian animals Late Devonian genus extinctions Fossil taxa described in 1839 Paleozoic life of Ontario Floresta Formation Paleozoic life of New Brunswick Paleozoic life of the Northwest Territories Paleozoic life of Quebec Phacopida genera