Thylacocephala

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Thylacocephala
Temporal range: Telychian to Late Cretaceous (possible early Cambrian record)
Thylacares brandonensis S12862-014-0159-2-10.jpg
Reconstruction of Thylacares , the earliest confirmed Thylacocephalan
Clausocaris lithographica reconstruction.png
Restoraton of Clausocaris , a Concavicaridan
Scientific classification
Kingdom:
Phylum:
Subphylum:
Class:
Thylacocephala

Pinna et al., 1982
Orders

The Thylacocephala (from the Greek θύλακος or thylakos, meaning "pouch", and κεφαλή or cephalon meaning "head") are a unique group of extinct arthropods, with possible crustacean affinities. As a class they have a short research history, having been erected in the early 1980s. [1] [2] [3]

Contents

They typically possess a large, laterally flattened carapace that encompasses the entire body. The compound eyes tend to be large and bulbous, and occupy a frontal notch on the carapace. They possess three pairs of large raptorial limbs, and the abdomen bears a battery of small swimming limbs.

The earliest thylacocephalan fossil is thought to date from the lower Cambrian, [4] while the class has a definite presence in Lower Silurian marine communities. [5] As a group, the Thylacocephala survived to the Upper Cretaceous. [6]

Beyond this, there remains much uncertainty concerning fundamental aspects of the thylacocephalan anatomy, mode of life, and relationship to the Crustacea, with whom they have always been cautiously aligned.

Research history

Fossil of Triassic Ankitokazocaris Ankitokazocaris.JPG
Fossil of Triassic Ankitokazocaris

The Thylacocephala is only recently described as a class, yet species now included within the group were first described at the turn of the century. [7] [8] These were typically assigned to the phyllocarids despite an apparent lack of abdomen and appendages. [9] In 1982/83, three research groups independently created higher taxa to accommodate new species. Based on a specimen from northern Italy, Pinna et al. designated a new class, [1] Thylacocephala, while Secrétan – studying Dollocaris ingens, a species from the La Voulte-sur-Rhône konservat-lagerstätte in France – erected the class Conchyliocarida. [10] Briggs & Rolfe, working on fossils from Australia's Devonian deposits were unable to attribute certain specimens to a known group, and created an order of uncertain affinities, the Concavicarida, to accommodate them. [3] It was apparent the three groups were in fact working on a single major taxon (Rolfe noted disagreements over interpretation and taxonomic placement largely resulted from a disparity of sizes and differences in preservation.) [11] The group took the name Thylacocephala by priority, with Concavicarida and Conchyliocarida subjugated to orders, erected by Rolfe, [11] and modified by Schram. [9]

Taxonomy

Ostenocaris cypriformis Ostenocaris cypriformis.JPG
Ostenocaris cypriformis

Researchers agree the Thylacocephala represent a class. Some efforts have been made at further classification: Schram split currently known taxa into two orders: [9]

The accuracy of this scheme has been questioned in recent papers, [4] as it stresses differences in the eyes and exoskeletal structure, which – in modern arthropods – tend to be a response to environmental conditions. Thus it has been suggested these features are too strongly controlled by external factors to be used alone to distinguish higher taxa. The problem is exacerbated by the limited number of thylacocephalan species known. More reliable anatomical indicators would include segmentation and appendage attachments (requiring the internal anatomy, currently elusive as a result of the carapace).

Anatomy

Based on Vannier, [4] modified after Schram: [9] The Thylacocephala are bivalved arthropods with morphology exemplified by three pairs of long raptorial (predatory) appendages and hypertrophied. They have a worldwide distribution. A laterally compressed, shield−like carapace encloses the entire body, and often has an anterior rostrum−notch complex and posterior rostrum. Its lateral surface can be externally ornamented, and evenly convex or with longitudinal ridges. Spherical or drop-shaped eyes are situated in the optic notches, and are often hypertrophied, filling the notches or forming a paired, frontal globular structure. No prominent abdominal features emerge from the carapace, and the cephalon is obscured. Even so, some authors have suggested the presence of five cephalic appendages, three of which could be the very long genticulate and chelate raptorials protruding beyond the ventral margin. [9] [12] Alternatively these could originate from three anterior trunk segments. [13] The posterior trunk has a series of eight to twenty [3] [5] styliform, filamentous pleopod-like appendages, decreasing in size posteriorly. Most Thylacocephala have eight pairs of well developed gills, found in the trunk region.

Beyond this there is a lack of knowledge about even basic thylacocephalan anatomy, including the number of posterior segments, origin of the raptorials, number of cephalic appendages, shape and attachment of gills, character of mouth, stomach and gut. This results from the class's all–encompassing carapace, which prevents the study of their internal anatomy in fossils.

Affinities

It is universally accepted that the Thylacocephala are arthropods, yet the position within this phylum is debated. It has always been cautiously assumed that the class is a member of the Crustacea, but no conclusive proof exists. The strongest apomorphy aligning the class with other crustaceans is the carapace. As this feature has evolved independently numerous times within the Crustacea and other arthropods, it is not a very reliable pointer, and such evidence alone remains insufficient to align the class with the crustaceans. [9]

Of the features which could prove crustacean affinities, the arrangement of mouthparts would be the easiest to find in the Thylacocephala. The literature features some mention of such a head arrangement, but none definitive. Schram reports the discovery of mandibles in the Mazon Creek thylacocephalan Concavicaris georgeorum. [9] Secrétan also mentions – with caution – possible mandibles in serial sections of Dollocaris ingens, and traces of small limbs in the cephalic region (not well preserved enough to assess their identity). [13] Lange et al. report a new genus and species, Thylacocephalus cymolopos, from the Upper Cretaceous of Lebanon, which has two possible pairs of antennae, but note the possession of two pairs of antennae alone does not prove the class occupies a position in the crown-group Crustacea. [14]

Despite a lack of evidence for a crustacean body plan, several authors have aligned the class with different groups of crustaceans. Schram provides an overview of possible affinities: [9]

In these various interpretations, numerous different limb arrangements for the three raptorials have been proposed:

Further work is necessary to provide any solid conclusions.

Disagreements

Numerous conflicts of opinion surround the Thylacocephala, of which the split between the “Italian school” and rest of the world is the most notable. Based on poorly preserved fossils from the Osteno deposits of Lombardy, Pinna et al. erected the class Thylacocephala. [12] Based on inferred cirripede affinities the authors concluded the frontal lobed structure was not an eye, but a 'cephalic sac'. This opinion arose from the misinterpretation of the stomach as a reproductive organ (its contents included vertebral elements of fish, thought to be ovarian eggs). [4] Such an arrangement is reminiscent of cirripede crustaceans, leading the authors to suggest a sessile, filter feeding mode of life, the 'cephalic sac' used to anchor the organism to the seabed. The researchers have since conceded it is highly improbable the ovaries are situated in the head, but maintain that the frontal structure is not an eye. Instead they suggest the 'cephalic sac' is covered with microsclerites, their arguments most recently presented in Alessandrello et al. [16]

Instead the authors suggest the sac is used to break down coarse chunks of food and reject indigestible portions.

All other parties interpret this as a large compound eye, the hexagons being preserved ommatidia (all researchers agree these are the same structure). [11] [13] This is supported by fossils of Dollocaris ingens which are so well preserved that individual retinula cells can be discerned. The preservation is so exceptional that studies have shown the species' numerous small ommatidia, distributed over the large eyes, could reduce the angle between ommatidia, thus improve their ability to detect small objects. [17] Of the arguments above, it is posited by opponents that eyes are complex structures, and those in the Thylacocephala display clear and numerous affinities with compound eyes in other arthropod fossils, down to a cellular level of detail. The 'cephalic sac' structure itself is poorly preserved in Osteno specimens, a possible reason for interstitial 'sclerites'. The structural analogy with a cirripede peduncle lost supporting evidence when the 'ovaries' were shown to be alimentary residues, [4] and the sac muscular system could be used to support the eyes. The unusual position of the stomach is thus the strongest inconsistency, but the Thylacocephala are defined by their unusual features, so this is not inconceivable. Further, Rolfe suggests the eyes' position can be explained if they have a large posterior area of attachment, [11] while Schram suggests that the stomach region extending into the cephalic sac could result from an inflated foregut or anteriorly directed caecum. [9]

Discussion of the matter has ceased in the last decade, and most researchers accept the anterior structure is an eye. Confusion is most likely the result of differing preservation in Osteno.

Mode of life

Numerous modes of life have been suggested for the Thylacocephala.

Secrétan suggested Dollocaris ingens was too large to swim, [13] so inferred a predatory 'lurking' mode of life, lying in wait on the sea bed and then springing out to capture prey. The author also suggested it could be necrophagous, supported by Alessandrello et al., [16] who suggest they would have been incapable of directly killing the shark remains found in the Osteno specimens' alimentary residues. Instead they surmise the Thylacocephala could have ingested shark vomit which included such remains.

Vannier et al. note the Thylacocephala possess features which would suggest adaptations for swimming in dim-light environments – a thin, non-mineralized carapace, well-developed rostral spines for possible buoyancy control in some species, a battery of pleopods for swimming, and large prominent eyes. [4] This is supported by the Cretaceous species from Lebanon, which show adaptations for swimming, [6] and possibly schooling.

Rolfe provides many possibilities, but concludes a realistic mode of life is mesopelagic, by analogy with hyperiid amphipods. [11] Further suggests floor-dwelling is also possible, and that the organism could rise to catch prey during the day and return to the sea floor at night. Another notable proposal is that, like hyperiids, the class could gain oil from their food source for buoyancy, an idea supported by their diet (known from stomach residues containing shark and coleoid remains, and other Thylacocephala).

Alessandrello et al. suggest a head-down, semi-sessile life on a soft bottom, [16] in agreement with that of Pinna et al., based on cirripede affinities. A necrophagous diet is suggested. [12]

Briggs & Rolfe report that all the Gogo Thylacocephala are found in a reef formation, suggesting a shallow water environment. [3] The authors speculate that due to the terracing of the carapace an infaunal mode of life is possible, or the ridges could provide more friction for hiding in crevices of rock.

Schram suggests a dichotomy in size of the class results from different environments; [9] larger Thylacocephala could have lived in a fluid characterized by turbulent flow, and relied on single power stroke of trunk limbs to position themselves. He suggests that smaller forms may have resided in a viscous medium, characterized by laminar flow, and used a lever to generate the speed necessary to capture prey.

Related Research Articles

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Agnostida is an order of arthropod which first developed near the end of the Early Cambrian period and thrived during the Middle Cambrian. They are present in the Lower Cambrian fossil record along with trilobites from the Redlichiida, Corynexochida, and Ptychopariida orders. The last agnostids went extinct in the Late Ordovician.

Trilobite Class of extinct, Paleozoic arthropods

Trilobites are a group of extinct marine artiopodan 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 252 million years ago. Trilobites were among the most successful of all early animals, existing in oceans for almost 300 million years.

Malacostraca Largest class of crustaceans

Malacostraca is the largest of the six classes of crustaceans, containing about 40,000 living species, divided among 16 orders. Its members, the malacostracans, display a great diversity of body forms and include crabs, lobsters, crayfish, shrimp, krill, woodlice, amphipods, mantis shrimp and many other, less familiar animals. They are abundant in all marine environments and have colonised freshwater and terrestrial habitats. They are segmented animals, united by a common body plan comprising 20 body segments, and divided into a head, thorax, and abdomen.

Remipedia

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Carapace Part of exoskeleton in some animals

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 underside is called the plastron.

Decapod anatomy The entire structure of a decapod crustacean

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Eucarida

Eucarida is a superorder of the Malacostraca, a class of the crustacean subphylum, comprising the decapods, krill, Amphionides and Angustidontida. They are characterised by having the carapace fused to all thoracic segments, and by the possession of stalked eyes.

Cheloniellida Order of arthropods (fossil)

Cheloniellida is a taxon of extinct Paleozoic arthropods. As of 2018, 8 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.

<i>Waptia</i> Species of crustacean (fossil)

Waptia fieldensis is an extinct species of arthropod from the Middle Cambrian Burgess Shale Lagerstätte of Canada. It grew to a length of about 8 cm (3 in) and resembled modern shrimp in both morphology and habit. It had a large bivalved carapace and a segmented body terminating into a pair of tail flaps. It was an active swimmer, feeding on organic particles it gathered from the seafloor substrate. It is also one of the oldest animals with direct evidence of brood care.

Edriophthalma is a disused peracarid (Malacostraca) classification comprising Isopoda and Amphipoda, first proposed by William Elford Leach in 1815. They have several common features, such as the fact that they both lack a carapace, possess sessile compound eyes, and thoracic coxae fused to their pleurites. Some molecular studies have shown that these are not related. The group has also been known as Acaridea and Arthrostraca.

<i>Angustidontus</i>

Angustidontus is a genus of predatory pelagic crustaceans from the Late Devonian and Early Carboniferous periods, classified as part of the subclass Eumalacostraca. Fossils of the genus have been recovered in relative abundance from Canada, Germany, the Czech Republic, Poland, Belarus, Ukraine and large parts of the United States, including Oklahoma, Ohio, Indiana, Kentucky, Montana, Utah, Nevada.

Radiodonta

Radiodonta is an order of stem-group arthropods that was successful worldwide during the Cambrian period, and included the earliest large predators known. They may be referred to as radiodonts, radiodontans, radiodontids, anomalocarids, or anomalocaridids, although the latter originally refer to the family Anomalocarididae, which previously included all species of this order but is now restricted to only a few species. Some of the most famous species of radiodonts are the Cambrian taxa Anomalocaris canadensis, Hurdia victoria, Peytoia nathorsti, and Amplectobelua symbrachiata, the Ordovician Aegirocassis benmoulai and the Devonian Schinderhannes bartelsi.

Phylogeny of Malacostraca is the evolutionary relationships of the largest of the six classes of crustaceans, containing about 40,000 living species, divided among 16 orders. Its members display a great diversity of body forms. Although the class Malacostraca is united by a number of well-defined and documented features, which were recognised a century ago by William Thomas Calman in 1904, the phylogenetic relationship of the orders which compose this class is unclear due to the vast diversity present in their morphology. Molecular studies have attempted to infer the phylogeny of this clade, resulting in phylogenies which have a limited amount of morphological support. To resolve a well-supported eumalacostracan phylogeny and obtain a robust tree, it will be necessary to look beyond the most commonly utilized sources of data.

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".

<i>Lepidocaris</i>

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Crustacean Subphylum of arthropods

Crustaceans form a large, diverse arthropod taxon which includes such animals as crabs, lobsters, crayfish, shrimps, prawns, krill, woodlice, and barnacles. The crustacean group can be treated as a subphylum under the clade Mandibulata; because of recent molecular studies it is now well accepted that the crustacean group is paraphyletic, and comprises all animals in the clade Pancrustacea other than hexapods. Some crustaceans are more closely related to insects and the other hexapods than they are to certain other crustaceans.

<i>Ostenocaris</i>

Ostenocaris is a Jurassic species of giant Thylacocephalan crustacean, sufficiently distinct from its relatives to be placed in its own family, Ostenocarididae. It is believed to be a bethonic animal and one of the most important necrophagus animals of its environment.

<i>Clausocaris</i>

Clausocaris is an extinct genus of Thylacocephalan containing the single species Clausocaris lithographica from the Late Jurassic (Tithonian) aged Solnhofen Limestone in Germany. It was originally named Clausia by Oppenheim in 1888, but was later changed to Clausocaris. The morphology suggests a lifestyle of a mobile or ambush oceanic predator.

Moltrasio Formation

The Moltrasio Formation also known as Lombardische Kieselkalk Formation is a geological formation in Italy. This Formation mostly developed on the Lower ot Middle Sinemurian stage of the Lower Jurassic, where on the Lombardian basin tectonic activity modified the current marine and terrestrial habitats. Here it was developed a series of marine-related depositional settings, represented by an outcrop of 550–600 m of grey Calcarenites and Calcilutites with chert lenses and marly interbeds, that recovers the Sedrina, Moltrasio and Domaro Formations. This was mostly due to the post-Triassic crisis, that was linked locally to tectonics. The Moltrasio Formation is considered a continuation of the Sedrina Limestone and the Hettangian Albenza Formation, and was probably a shallow water succession, developed on the pasive margin of the westernmost Southern Alps. It is known due to the exquisite preservation observed on the Outcrop on Osteno, where several kinds of marine biota have been recovered.

<i>Dollocaris</i>

Dollocaris was a genus of Thylacocephalan that lived during the Jurassic period. Fossils have been found in France, specifically the La Voulte-sur-Rhône lagerstätte. It is known for its massive compound eyes, giving Dollocaris a rather characteristic appearance. One species is currently known, D. ingens.

References

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