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The cephalon is the head section of an arthropod. It is a tagma, i.e., a specialized grouping of arthropod segments. The word cephalon derives from the Greek κεφαλή (kephalē), meaning "head".
In insects, head is a preferred term. The insect head consists of five segments, including three (the labial, maxillary and mandibular) necessary for food uptake, which are altogether known as the gnathocephalon and house the suboesophageal ganglion of the brain, as well as the antennal segment, and an ocular segment, as well as a non segmented fused section of the head where the archicerebrum is housed known as the acron. [1] See also arthropod head problem.
In chelicerates and crustaceans, the cephalothorax is derived from the fusion of the cephalon and the thorax, and is usually covered by a single unsegmented carapace. In relation with the arthropod head problem, phylogeny studies show that members of the Malacostraca class of crustaceans have five segments in the cephalon, when not fused with the thorax to form a cephalothorax.
In the Late Precambrian or Lower Cambrian Proarticulata species Praecambridium sigillum , that superficially resembles a trilobite, the term is also used to describe the anterior part of the animal.
The head of the Thylacocephala is also referred to as a cephalon: the cephalon is usually obscured by the carapace. Thylacocephala are a unique group of extinct arthropods, with possible crustacean affinities, thought to occur from the lower Cambrian, but with certainty between the Lower Silurian and the Upper Cretaceous.
The cephalon of trilobites is highly variable with a lot of morphological complexity. The glabella, the expression of the axial lobe in the cephalon, forms a dome underneath which sat the "crop" or "stomach". Generally the exoskeleton has few distinguishing ventral features, but the cephalon often preserves muscle attachment scars and occasionally the hypostome, a small rigid plate comparable to the ventral plate in other arthropods. A toothless mouth and stomach sat upon the hypostome with the mouth facing backwards at the rear edge of the hypostome.
Hypostome morphology is highly variable; sometimes supported by an un-mineralised membrane (natant), sometimes fused onto the anterior doublure with an outline very similar to the glabella above (conterminant) or fused to the anterior doublure with an outline significantly different from the glabella (impendent). Many variations in shape and placement of the hypostome have been described. [2] The size of the glabella and the lateral fringe of the cephalon, together with hypostome variation, have been linked to different lifestyles, diets and specific ecological niches. [3]
The lateral fringe of the cephalon is greatly exaggerated in the Harpetida, in other species a bulge in the pre-glabellar area is preserved that suggests a brood pouch. [4] Highly complex compound eyes are another obvious feature of the cephalon.
When trilobites moulted, the librigenae ("free cheeks") separated along the facial suture to assist moulting, leaving the cranidium (glabella + fixigenae) exposed. Trilobite facial sutures can be roughly divided into three main types (proparian, gonatoparian, and opisthoparian) according to where the sutures end relative to the genal angle (the edges where the side and rear margins of the cephalon converge). Early Cambrian trilobites belonging to the suborder Olenellina (like Fallotaspis ) lacked facial sutures. Other later trilobites also lost facial sutures secondarily. [5]
Agnostida are an order of extinct arthropods which have classically been seen as a group of highly modified trilobites, though some recent research has doubted this placement. Regardless, they appear to be close relatives as part of the Artiopoda. They are present in the Lower Cambrian fossil record along with trilobites from the Redlichiida, Corynexochida, and Ptychopariida orders, and were highly diverse throughout the Cambrian. Agnostidan diversity severely declined during the Cambrian-Ordovician transition, and the last agnostidans went extinct in the Late Ordovician.
Trilobites 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 Atdabanian stage of the Early Cambrian period and they flourished throughout the lower Paleozoic before slipping into a long decline, when, during the Devonian, all trilobite orders except the Proetida died out. The last extant trilobites finally disappeared in the mass extinction at the end of the Permian about 251.9 million years ago. Trilobites were among the most successful of all early animals, existing in oceans for almost 270 million years, with over 22,000 species having been described.
Redlichiida is an order of trilobites, a group of extinct marine arthropods. Species assigned to the order Redlichiida are among the first trilobites to appear in the fossil record, about halfway during the Lower Cambrian. Due to the difficulty to relate sediments in different areas, there remains some discussion, but among the earliest are Fallotaspis, and Lemdadella, both belonging to this order. The first representatives of the orders Corynexochida and Ptychopariida also appear very early on and may prove to be even earlier than any redlichiid species. In terms of anatomical comparison, the earliest redlichiid species are probably ancestral to all other trilobite orders and share many primitive characters. The last redlichiid trilobites died out before the end of the Middle Cambrian.
The cephalothorax, also called prosoma in some groups, is a tagma of various arthropods, comprising the head and the thorax fused together, as distinct from the abdomen behind.. The word cephalothorax is derived from the Greek words for head and thorax. This fusion of the head and thorax is seen in chelicerates and crustaceans; in other groups, such as the Hexapoda, the head remains free of the thorax. In horseshoe crabs and many crustaceans, a hard shell called the carapace covers the cephalothorax.
Dalmanites is a genus of trilobite in the order Phacopida. They lived from the Late Ordovician to Middle Devonian.
Paradoxides is a genus of large to very large trilobite found throughout the world during the Middle Cambrian period. One record-breaking specimen of Paradoxides davidis, described by John William Salter in 1863, is 37 cm (15 in). The cephalon was semicircular with free cheeks ending in long, narrow, recurved spines. Eyes were crescent shaped providing an almost 360° view, but only in the horizontal plane. Its elongate thorax was composed of 19–21 segments and adorned with longish, recurved pleural spines. Its pygidium was comparatively small. Paradoxides is a characteristic Middle-Cambrian trilobite of the 'Atlantic' (Avalonian) fauna. Avalonian rocks were deposited near a small continent called Avalonia in the Paleozoic Iapetus Ocean. Avalonian beds are now in a narrow strip along the East Coast of North America, and in Europe.
Emuellidae are a small family of trilobites, a group of extinct marine arthropods, that lived during the late Lower Cambrian of the East Gondwana supercontinent, in what are today South-Australia and Antarctica.
Redlichiina is a suborder of the order Redlichiida of Trilobites. The suborder contains three superfamilies: Emuelloidea, Redlichioidea and Paradoxidoidea. These trilobites are some of the oldest trilobites known. They originated at the beginning of the Cambrian Period and disappeared at the end of the middle Cambrian.
In biology, a tagma is a specialized grouping of multiple segments or metameres into a coherently functional morphological unit. Familiar examples are the head, the thorax, and the abdomen of insects. The segments within a tagma may be either fused or so jointed as to be independently moveable.
Cheloniellida is a taxon of extinct Paleozoic arthropods. As of 2018, 7 monotypic genera of cheloniellids had been formally described, whose fossils are found in marine strata ranging from Ordovician to Devonian in age. Cheloniellida has a controversial phylogenetic position, with previous studies associated it as either a member or relative of various fossil and extant arthropod taxa. It was later accepted as a member of Vicissicaudata within Artiopoda.
The (pan)arthropod head problem is a long-standing zoological dispute concerning the segmental composition of the heads of the various arthropod groups, and how they are evolutionarily related to each other. While the dispute has historically centered on the exact make-up of the insect head, it has been widened to include other living arthropods, such as chelicerates, myriapods, and crustaceans, as well as fossil forms, such as the many arthropods known from exceptionally preserved Cambrian faunas. While the topic has classically been based on insect embryology, in recent years a great deal of developmental molecular data has become available. Dozens of more or less distinct solutions to the problem, dating back to at least 1897, have been published, including several in the 2000s.
Dikelocephalus is a genus of very large trilobites of up to 50 cm (20 in) long, that lived during the last 3 million years of the Cambrian (Sunwaptan). Their fossils are commonly found as disarticulated sclerites, in the upper Mississippi Valley and in Canada (Alberta). The exoskeleton is rounded anteriorly, with the thorax and sides of the tailshield slightly tapering to about 2⁄3× of the width across the base of the spines at the back of the headshield. At the side corners of the pygidium there may be triangular or hooked spines, pointing backwards, while between the spines the posterior margin is at a 30-75° angle with the lateral margin, gently convex or nearly straight. If pygidial spines are lacking, the margin is gradually rounded. The thorax has 12 segments.
Tsunyidiscus is a trilobite belonging to the Suborder Eodiscina. Tsunyidiscus appeared near the end of the Lower Cambrian, during the late Atdabanian stage of geologic time and some collections suggest it may have survived into the Botomian. The genus is very small, oculate and isopypous with a narrow dome-shaped glabella and a narrow bullet-shaped pygidial axis. Thorax consists of three segments. Tsunyidiscus is the only genus currently attributed to the family Tsunyidiscidae.
Phytophilaspis is a phosphatized genus of trilobite-like arthropod with eyes, found in association with algal remains. It dwelt in well-lit, shallow waters.
The hypostome is the hard mouthpart of trilobites found on the ventral side of the cephalon (head). The hypostome can be classified into three types based on whether they are permanently attached to the rostrum or not and whether they are aligned to the anterior dorsal tip of the glabella.
Conocoryphe is a genus of primarily eyeless trilobites belonging to the family Conocoryphidae. They lived during the Middle Cambrian period, about 505 million years ago. These arthropods lived on the sea bottom (epifaunal) and lived off dead particulate organic matter.
Odontochile is a genus of trilobites in the order Phacopida, family Dalmanitidae.
Eodiscina is trilobite suborder. The Eodiscina first developed near the end of the Lower Cambrian period and became extinct at the end of the Middle Cambrian. Species are tiny to small, and have a thorax of two or three segments. Eodiscina includes six families classified under one superfamily, Eodiscoidea.
Orygmaspis is a genus of asaphid trilobite with an inverted egg-shaped outline, a wide headshield, small eyes, long genal spines, 12 spined thorax segments and a small, short tailshield, with four pairs of spines. It lived during the Upper Cambrian in what are today Canada and the United States.
Viaphacops is a genus of trilobites in the order Phacopida, family Phacopidae, that lived during the Middle Devonian, and is known from North and South America, Asia.