[23] Principal evolutionary trends from primitive morphologies, such as exemplified by Eoredlichia,[24] include the origin of new types of eyes, improvement of enrollment and articulation mechanisms, increased size of pygidium (micropygy to isopygy), and development of extreme spinosity in certain groups.

[24] Specific changes to the cephalon are also noted; variable glabella size and shape, position of eyes and facial sutures, and hypostome specialization.

[15] Very shortly after trilobite fossils appeared in the lower Cambrian, they rapidly diversified into the major orders that typified the Cambrian—Redlichiida, Ptychopariida, Agnostida, and Corynexochida.

[23] A continuing decrease in Laurentian continental shelf area[28] is recorded at the same time as the extinctions, suggesting major environmental upheaval.

Decreasing diversity[32] of genera limited to shallow-water shelf habitats coupled with a drastic lowering of sea level (regression) meant that the final decline of trilobites happened shortly before the end Permian mass extinction event.

Because they appeared quickly in geological time, and moulted like other arthropods, trilobites serve as excellent index fossils, enabling geologists to date the age of the rocks in which they are found.

[39] The fossils are dated to the Givetian (387.2–382.7 million years ago) when the Western New York Region was 30 degrees south of the equator and completely covered in water.

[38] In 1994, the quarry became Penn Dixie Fossil Park & Nature Reserve when they received 501(c)3 status and was opened for visitation and collection of trilobite samples.

[41] A famous location for trilobite fossils in the United Kingdom is Wren's Nest, Dudley, in the West Midlands, where Calymene blumenbachii is found in the Silurian Wenlock Group.

The French palaeontologist Joachim Barrande (1799–1883) carried out his landmark study of trilobites in the Cambrian, Ordovician and Silurian of Bohemia, publishing the first volume of Système silurien du centre de la Bohême in 1852.

The study of Paleozoic trilobites in the Welsh-English borders by Niles Eldredge was fundamental in formulating and testing punctuated equilibrium as a mechanism of evolution.

The 10 most commonly recognized trilobite orders are Agnostida, Redlichiida, Corynexochida, Lichida, Odontopleurida, Phacopida, Proetida, Asaphida, Harpetida and Ptychopariida.

[28] However, a partial specimen of the Ordovician trilobite Hungioides bohemicus found in 2009 in Arouca, Portugal is estimated to have measured when complete 86.5 cm (34.1 in) in length.

Despite morphological complexity and an unclear position within higher classifications, there are a number of characteristics which distinguish the trilobites from other arthropods: a generally sub-elliptical, dorsal, chitinous exoskeleton divided longitudinally into three distinct lobes (from which the group gets its name); having a distinct, relatively large head shield (cephalon) articulating axially with a thorax comprising articulated transverse segments, the hindmost of which are almost invariably fused to form a tail shield (pygidium).

[61] 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.

[8] The anterior and lateral fringe of the cephalon is greatly enlarged in the Harpetida, in other species a bulge in the pre-glabellar area is preserved that suggests a brood pouch.

[73] All species assigned to the suborder Olenellina, that became extinct at the very end of the Early Cambrian (like Fallotaspis, Nevadia, Judomia, and Olenellus) lacked facial sutures.

The course of the facial sutures from the front of the visual surface varies at least as strongly as it does in the rear, but the lack of a clear reference point similar to the genal angle makes it difficult to categorize.

This is known in Triarthrus, and in the Phacopidae, but in that family the facial sutures are not functional, as can be concluded from the fact that free cheeks are not found separated from the cranidium.

[83] Even in an agnostid, with only 2 articulating thoracic segments, the process of enrollment required a complex musculature to contract the exoskeleton and return to the flat condition.

The exact purpose of the prosopon is not resolved but suggestions include structural strengthening, sensory pits or hairs, preventing predator attacks and maintaining aeration while enrolled.

[97] Improving eyesight of both predator and prey in marine environments has been suggested as one of the evolutionary pressures furthering an apparent rapid development of new life forms during what is known as the Cambrian explosion.

[23] The fossil record of trilobite eyes is complete enough that their evolution can be studied through time, which compensates to some extent for the lack of preservation of soft internal parts.

[101] Rigid calcite lenses would have been unable to accommodate to a change of focus like the soft lens in a human eye would; in some trilobites, the calcite formed an internal doublet structure,[102] giving superb depth of field and minimal spherical aberration, according to optical principles discovered by French scientist René Descartes and Dutch physicist Christiaan Huygens in the 17th century.

In Proetida and Phacopina from western Europe and particularly Tropidocoryphinae from France (where there is good stratigraphic control), there are well studied trends showing progressive eye reduction between closely related species that eventually leads to blindness.

[111] Despite the absence of supporting fossil evidence, their similarity to living arthropods has led to the belief that trilobites multiplied sexually and produced eggs.

[115] Pelagic larval life-style proved ill-adapted to the rapid onset of global climatic cooling and loss of tropical shelf habitats during the Ordovician.

Edward Lhwyd published in 1698 in The Philosophical Transactions of the Royal Society, the oldest scientific journal in the English language, part of his letter "Concerning Several Regularly Figured Stones Lately Found by Him", that was accompanied by a page of etchings of fossils.

[118] One of his etchings depicted a trilobite he found near Llandeilo, probably on the grounds of Lord Dynefor's castle, he described as "the skeleton of some flat Fish".

He proposed to call the Dudley specimens Pediculus marinus major trilobos (large trilobed marine louse), a name which lasted well into the 19th century.