Cephalon (trilobite)
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The cephalon is the
head A head is the part of an organism which usually includes the ears, brain, forehead, cheeks, chin, eyes, nose, and mouth, each of which aid in various sensory functions such as sight, hearing, smell, and taste. Some very simple ani ...
section of an
arthropod Arthropods ( ) are invertebrates in the phylum Arthropoda. They possess an arthropod exoskeleton, exoskeleton with a cuticle made of chitin, often Mineralization (biology), mineralised with calcium carbonate, a body with differentiated (Metam ...
. It is a tagma, i.e., a specialized grouping of arthropod segments. The word cephalon derives from the Greek κεφαλή (kephalē), meaning "head".


Insects

In
insect Insects (from Latin ') are Hexapoda, hexapod invertebrates of the class (biology), class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (Insect morphology#Head, head, ...
s, ''
head A head is the part of an organism which usually includes the ears, brain, forehead, cheeks, chin, eyes, nose, and mouth, each of which aid in various sensory functions such as sight, hearing, smell, and taste. Some very simple ani ...
'' 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 The suboesophageal ganglion (acronym: SOG; synonym: ''subesophageal ganglion'') of arthropods and in particular insects is part of the arthropod central nervous system (CNS). As indicated by its name, it is located ''below the'' ''oesophagus'', insi ...
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. See also
arthropod head problem The (pan)arthropod head problem is a long-standing zoological dispute concerning the Segmentation (biology), segmental composition of the heads of the various arthropod groups, and how they are evolutionarily related to each other. While the dis ...
.


Chelicerates and crustaceans

File:Cherax warsamsonicus - ZooKeys 660 151-167 (cropped).jpg, The crustacean '' Cherax warsamsonicus'' File:Phrynus asperatipes.jpg, The amplypygid chelicerate '' Phrynus asperatipes'' In
chelicerate The subphylum Chelicerata (from Neo-Latin, , ) constitutes one of the major subdivisions of the phylum Arthropoda. Chelicerates include the sea spiders, horseshoe crabs, and arachnids (including harvestmen, scorpions, spiders, solifuges, ticks ...
s and
crustacean 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 ...
s, the
cephalothorax 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 terms ''prosoma'' and ''opisthosoma'' are equivalent to ''cepha ...
is derived from the fusion of the cephalon and the thorax, and is usually covered by a single unsegmented
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 unde ...
. In relation with the
arthropod head problem The (pan)arthropod head problem is a long-standing zoological dispute concerning the Segmentation (biology), segmental composition of the heads of the various arthropod groups, and how they are evolutionarily related to each other. While the dis ...
,
phylogeny A phylogenetic tree or phylogeny is a graphical representation which shows the evolutionary history between a set of species or Taxon, taxa during a specific time.Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, M ...
studies show that members of the
Malacostraca Malacostraca is the second largest of the six classes of pancrustaceans behind insects, containing about 40,000 living species, divided among 16 orders. Its members, the malacostracans, display a great diversity of body forms and include crab ...
class of crustaceans have five segments in the cephalon, when not fused with the thorax to form a cephalothorax.


Proarticulata

In the Late Precambrian or Lower Cambrian
Proarticulata Proarticulata is a proposed phylum of extinct, near-bilaterally symmetrical animals known from fossils found in the Ediacaran (Vendian) marine deposits, and dates to approximately . The name comes from the Greek () = "before" and Articulata, i. ...
species ''
Praecambridium sigillum ''Praecambridium sigillum'' is an extinct organism that superficially resembles a segmented trilobite-like arthropod. It was originally described as being a trilobite-like arthropod, though the majority of experts now place it within the Proartic ...
'', that superficially resembles a trilobite, the term is also used to describe the anterior part of the animal.


Thylacocephala

The head of the
Thylacocephala The Thylacocephala (from the Greek or ', meaning " pouch", and or ' meaning "head") are group of extinct probable mandibulate arthropods, that have been considered by some researchers as having possible crustacean affinities. As a class they ...
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 The Silurian ( ) is a geologic period and system spanning 23.5 million years from the end of the Ordovician Period, at million years ago ( Mya), to the beginning of the Devonian Period, Mya. The Silurian is the third and shortest period of t ...
and the Upper Cretaceous.


Trilobites

The cephalon of
trilobites 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, ...
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 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 ...
, 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. 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 niche In ecology, a niche is the match of a species to a specific environmental condition. Three variants of ecological niche are described by It describes how an organism or population responds to the distribution of Resource (biology), resources an ...
s. The lateral fringe of the cephalon is greatly exaggerated in the
Harpetida Harpetida is one of the eleven orders of the extinct arthropod class Trilobita. The first harpetid trilobites appear in the Upper Cambrian, and the last species die out in the late Devonian period. Harpetid trilobites are characterized among tr ...
, in other species a bulge in the pre-glabellar area is preserved that suggests a brood pouch. Highly complex compound eyes are another obvious feature of the cephalon.


Facial sutures

When trilobites moulted, the librigenae ("free cheeks") separated along the facial suture to assist moulting, leaving the cranidium (glabella + fixigenae) exposed.
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 ...
facial sutures can be roughly divided into three main types (
proparian 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, ...
, gonatoparian, and
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, ...
) 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 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 ...
trilobites belonging to the
suborder Order () is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between family and class. In biological classification, the order is a taxonomic rank used in the classification of organisms and recognized ...
Olenellina Olenellina is a suborder of the order Redlichiida of trilobites that occurs about halfway during the Lower Cambrian, at the start of the stage called the Atdabanian. Olenellina are arguably the earliest trilobites in the fossil record as members ...
(like ''
Fallotaspis ''Fallotaspis'' is a genus of redlichiid trilobites found in Early Cambrian-aged strata of the United States and Morocco Morocco, officially the Kingdom of Morocco, is a country in the Maghreb region of North Africa. It has coastlines o ...
'') lacked facial sutures. Other later trilobites also lost facial sutures secondarily.


See also

*
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 unde ...
*
Facial suture (trilobite) 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, ...
*
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 ...
*
Tagma (biology) In biology, a tagma (Greek: τάγμα, : tagmata – τάγματα - body of soldiers; battalion) is a specialized grouping of multiple segmentation (biology), segments or Metamerism (biology), metameres into a coherently functional Morphology ...
*
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 ...


References

{{reflist Arthropod anatomy Trilobite anatomy Animal head