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, existing in oceans for almost 270 million years, with over 22,000 species having been described. Because trilobites had wide diversity and an easily fossilized mineralised exoskeleton made of calcite, they left an extensive fossil record. The study of their fossils has facilitated important contributions to biostratigraphy, paleontology, evolutionary biology, and plate tectonics. Trilobites are placed within the clade Artiopoda, which includes many organisms that are morphologically similar to trilobites, but are largely unmineralised. The relationship of Artiopoda to other arthropods is uncertain. Trilobites evolved into many ecological niches; some moved over the seabed as predators, scavengers, or filter feeders, and some swam, feeding on plankton. Some even crawled onto land. Most lifestyles expected of modern marine arthropods are seen in trilobites, with the possible exception of parasitism (where scientific debate continues). Some trilobites (particularly the family Olenidae) are even thought to have evolved a symbiotic relationship with sulfur-eating bacteria from which they derived food. The largest trilobites were more than 70 centimetres (28 in) long and may have weighed as much as 4.5 kilograms (9.9 lb). The first appearance of trilobites in the fossil record defines the base of the Atdabanian/Cambrian Stage 3 time period of the Early Cambrian around 521 million years ago. Trilobites were already diverse and globally dispersed shortly after their origination, with trilobites reaching an apex of diversity during the late Cambrian–Ordovician, and remained diverse during the following Silurian and early Devonian. During the mid-late Devonian, their diversity strongly declined, being impacted by successive extinction events, including the Taghanic event, the Late Devonian mass extinction/Kellwasser event and the Hangenberg/end-Devonian mass extinction, wiping out most trilobite diversity and leaving Proetida as the only surviving order. Their diversity moderately recovered during the Early Carboniferous, before dropping to persistently low levels during the late Carboniferous and Permian periods, though they remained widespread until the end of their existence. The last trilobites disappeared in the end-Permian mass extinction event about 251.9 million years ago, by which time only a handful of species remained.
Evolution
Trilobite relatives Trilobites belong to the Artiopoda, a group of extinct arthropods morphologically similar to trilobites, though only the trilobites had heavily mineralised exoskeletons. Thus, other artiopodans are typically only found in exceptionally preserved deposits, mostly from the Cambrian period. The exact relationships of artiopods to other arthropods is uncertain. Some scholars consider them closely related to chelicerates (which include horseshoe crabs, sea spiders, and arachnids) as part of a clade called Arachnomorpha, while others consider them to be more closely related to Mandibulata (which contains insects, crustaceans and myriapods) as part of a clade called Antennulata.
Cladogram of Artiopoda including trilobites after Berks et al. 2023.
Fossil record of early trilobites
The earliest trilobites known from the fossil record are redlichiids and ptychopariid bigotinids dated to around 520 million years ago. Contenders for the earliest trilobites include Profallotaspis jakutensis (Siberia), Fritzaspis spp. (western US), Hupetina antiqua (Morocco) and Serrania gordaensis (Spain). Trilobites appeared at a roughly equivalent time in Laurentia, Siberia and West Gondwana. All Olenellina lack facial sutures (see below), and this is thought to represent the original state. The earliest sutured trilobite found so far (Lemdadella), occurs almost at the same time as the earliest Olenellina, suggesting the trilobites origin lies before the start of the Atdabanian, but without leaving fossils. Other groups show secondary lost facial sutures, such as all Agnostina and some Phacopina. Another common feature of the Olenellina also suggests this suborder to be the ancestral trilobite stock: early protaspid stages have not been found, supposedly because these were not calcified, and this also is supposed to represent the original state. Earlier trilobites may be found and could shed more light on their origins. Three specimens of a trilobite from Morocco, Megistaspis hammondi, dated 478 million years old contain fossilized soft parts. In 2024, researchers discovered soft tissues and other structures including the labrum in well-preserved trilobite specimens from Cambrian Stage 4 of Morocco, providing new anatomical information regarding the external and internal morphology of trilobites, and the cause of such extraordinary preservation is probably due to their rapid death after an underwater pyroclastic flow.
Divergence and extinction
Trilobites saw great diversification over time. For such a long-lasting group of animals, it is no surprise that trilobite evolutionary history is marked by a number of extinction events where some groups perished, and surviving groups diversified to fill ecological niches with comparable or unique adaptations. Generally, trilobites maintained high diversity levels throughout the Cambrian and Ordovician periods before entering a drawn-out decline in the Devonian, culminating in the final extinction of the last few survivors at the end of the Permian period.
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