Figures 1 and 3 | |||||||||
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Cladogram following von Reumont et al. (2012), figures 1 and 3. [13] |
Figure 2 | |||||||||
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Cladogram following von Reumont et al. (2012), figure 2. [13] |
Figure 4 | |||||||||
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Cladogram following von Reumont et al. (2012), figure 4. [13] |
Parts of this article (those related to documentation) need to be updated.(June 2022) |
Pancrustacea Temporal range: | |
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Sally Lightfoot crab ( Grapsus grapsus ). | |
Fruit fly ( Drosophila melanogaster ). | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Clade: | Mandibulata |
Clade: | Pancrustacea Zrzavý & Štys, 1997 |
Superclasses [1] and extinct orders | |
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Synonyms | |
TetraconataDohle, 2001 |
Pancrustacea is the clade that comprises all crustaceans and all hexapods (insects and relatives). [2] This grouping is contrary to the Atelocerata hypothesis, in which Hexapoda and Myriapoda are sister taxa, and Crustacea are only more distantly related. As of 2010, the Pancrustacea taxon was considered well accepted, with most studies recovering Hexapoda within Crustacea. [3] The clade has also been called Tetraconata, referring to having four cone cells in the ommatidia. [4] The term "Tetraconata" is preferred by some scientists in order to avoid confusion with the use of "pan-" to indicate a clade that includes a crown group and all of its stem group representatives. [5]
A monophyletic Pancrustacea has been supported by several molecular studies, [6] [7] [8] [9] [10] in most of which the subphylum Crustacea is paraphyletic with regard to hexapods (that is, that hexapods, including insects, are derived from crustacean ancestors). This means that within pancrustacea, only some members are actually crustaceans, hexapods being the main exception.
The evidence for this clade derives from molecular data and morphological characteristics. The molecular data consists of comparisons of nuclear ribosomal RNA genes, mitochondrial ribosomal RNA genes, and protein coding genes. The morphological data consists of ommatidial structures (see arthropod eye ), the presence of neuroblasts, and the form and style of axonogenesis by pioneer neurons. [11] [12]
In a 2005 study of nuclear genomes Regier et al. suggest that Hexapoda is most closely related to Branchiopoda and Cephalocarida + Remipedia, thereby hexapods are "terrestrial crustaceans", thus supporting the Pancrustacea hypothesis that maxillopods are not monophyletic (in the following cladograms Maxillopoda subclasses are highlighted). In addition, there appeared some evidence against the Ostracoda monophyly: that Ostracoda subclass Podocopa may form a clade with Branchiura. [6]
Cladogram following Regier et al. (2005). [6] |
A 2010 study of nuclear genomes (Regier et al.) strongly supports Pancrustacea and strongly favour Mandibulata (Myriapoda + Pancrustacea) over Paradoxopoda (Myriapoda + Chelicerata). According to this study, Pancrustacea is divided into four lineages: Oligostraca (Ostracoda, Mystacocarida, Branchiura, Pentastomida), Vericrustacea (Malacostraca, Thecostraca, Copepoda, Branchiopoda), Xenocarida (Cephalocarida, Remipedia) and Hexapoda, with Xenocarida as a sister group to the Hexapoda (comprising "Miracrustacea"). [7]
New clades proposed by Regier et al. are:
Of these proposed clades, only Multicrustacea was confirmed in later molecular studies.
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Cladogram following Regier et al. (2010). [7] |
In a 2012 molecular study, von Reumont et al. challenge the monophyly of Vericrustacea: they present four versions of Pancrustacea cladogram (figures 1–4), and in all four figures Remipedia is a sister group to Hexapoda, and Branchiopoda is a sister group to (Remipedia + Hexapoda). Thus, their data strongly suggest that Branchiopoda is more closely related to Hexapoda and Remipedia than to Multicrustacea. Based on these data, they propose the following scenario of evolution of Branchiopoda, Remipedia and Hexapoda: under the impact of predatory fishes their common ancestors go to the littoral zone, then ancestors of Branchiopoda go to the ephemeral freshwater habitat, whereas ancestors of Remipedia go to the anchialine cave, and ancestors of Hexapoda go to the land. [13]
Another molecular study (of mitochondrial genomes), conducted in 2012 by Jondeung et al., strongly support monophyletic Pancrustacea and places Malacostraca + Entomostraca and Branchiopoda as the sister clade to Hexapoda and places Cirripedia + Remipedia as a basal lineage of Pancrustacea. [14]
In 2013 combined study of morphology, including fossils, and molecular data, including expressed sequence tag, mitochondrial genome, nuclear genome, and ribosomal DNA data Oakley et al. obtained support for three pancrustacean clades: Oligostraca (Ostracoda, Mystacocarida, Branchiura, Pentastomida), Multicrustacea (Copepoda, Thecostraca, Malacostraca) and a clade they refer to as Allotriocarida (Branchiopoda, Cephalocarida, Remipedia, Hexapoda), as well as for monophyly of Ostracoda. Within Multicrustacea they obtained support for a clade they suggest the name Hexanauplia: Thecostraca + Copepoda. Relations within Allotriocarida remain uncertain: sister taxon to Hexapoda is either Remipedia, or the clade Branchiopoda + Cephalocarida, however, authors are inclined to the first version (see "Conclusion", 4), which is also consistent with von Reumont et al. (2012) results. [15] [13]
New proposed by Oakley et al. clades are:
Note: the Allotriocarida clade was also recovered in 2005 by Regier et al. as Clade #33, [6] but relations within it were different, and they did not choose a name for it.
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Cladogram following Oakley et al. (2013) [15] |
In 2013 Rota-Stabelli et al. used the signal in the 62 protein-coding genes assembled by Regier et al. in 2010 to improve the knowledge of the internal relationship in the Pancrustacea group. This data set infers a highly supported nucleotide tree that is substantially different from the corresponding, but poorly supported, amino acid one. The discrepancy between the nucleotide-based and the amino acids-based trees is caused by substitutions within synonymous codon families (especially those of serine-TCN and AGY): different arthropod lineages are differentially biased in their usage of serine, arginine, and leucine synonymous codons, and the serine bias is correlated with the topology derived from the nucleotides, but not the amino acids. The authors suggest that a parallel, partially compositionally driven, synonymous codon-usage bias affects the nucleotide topology. As substitutions between serine codon families can proceed through threonine or cysteine intermediates, amino acid data sets might also be affected by the serine codon-usage bias. The analyses suggests that a Dayhoff recoding strategy would partially ameliorate the effects of such bias. Although amino acids provide an alternative hypothesis of pancrustacean relationships, neither the nucleotides nor the amino acids version of this data set bring enough genuine phylogenetic information to robustly resolve the relationships within group, which should still be considered unresolved. However the amino acid tree seems to be more likely since it appears to be free from the synonymous codon-family bias affecting the nucleotide one. Most of the inferences based on amino acids sequences support a clade which includes Branchiopoda, Remipedia, Copepoda and Hexapoda (group A). Using the best amino acids substitution model, CATGTR, also Cephalocarida falls inside this group. In all the analyses group A (with or without Cephalocarida) is sister-group of a clade composed by Malacostraca, Oligostraca and Thecostraca (group B). [16]
The following image shows the tree resulting from the Dayhoff recoding.
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The relationship of Hexapoda and the crustacean classes is shown in the following phylogenetic tree, which shows Allotriocarida, along with Oligostraca and Multicrustacea, as the three main divisions of subphylum Pancrustacea, embracing the traditional crustaceans and the hexapods (including insects). [1]
According to Petrunina A.S. and Kolbasov G.A., the sixth subclass of Maxillopoda Tantulocarida may lie within Thecostraca, forming a clade with thecostracan infraclass Cirripedia (if so, Thecostraca excluding Tantulocarida is paraphyletic): [17] [18]
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Thecostraca cladogram following Petrunina (2012), page 19, picture 8 [18] |
Branchiopoda is a class of crustaceans. It comprises fairy shrimp, clam shrimp, Diplostraca, Notostraca, the Devonian Lepidocaris and possibly the Cambrian Rehbachiella. They are mostly small, freshwater animals that feed on plankton and detritus.
Tantulocarida is a highly specialised group of parasitic crustaceans that consists of about 33 species, treated as a class in superclass Multicrustacea. They are typically ectoparasites that infest copepods, isopods, tanaids, amphipods and ostracods.
Remipedia is a class of blind crustaceans, closely related to hexapods, found in coastal aquifers which contain saline groundwater, with populations identified in almost every ocean basin so far explored, including in Australia, the Caribbean Sea, and the Atlantic Ocean. The first described remipede was the fossil Tesnusocaris goldichi. Since 1979, at least seventeen living species have been identified in subtropical regions around the world.
Ostracods, or ostracodes, are a class of the Crustacea, sometimes known as seed shrimp. Some 33,000 species have been identified, grouped into 7 valid orders. They are small crustaceans, typically around 1 mm (0.04 in) in size, but varying from 0.2 to 30 mm in the case of the marine Gigantocypris. The largest known freshwater species is Megalocypris princeps, which reach 8 mm in length. In most cases, their bodies are flattened from side to side and protected by a bivalve-like valve or "shell" made of chitin, and often calcium carbonate. The family Entocytheridae and many planktonic forms do not have calcium carbonate. The hinge of the two valves is in the upper (dorsal) region of the body. Ostracods are grouped together based on shell and soft part morphology, and molecular studies have not unequivocally supported the group's monophyly. They have a wide range of diets, and the class includes carnivores, herbivores, scavengers and filter feeders, but most ostracods are deposit feeders.
Eucarida is a superorder of the Malacostraca, a class of the crustacean subphylum, comprising the decapods, krill, and Angustidontida. They are characterised by having the carapace fused to all thoracic segments, and by the possession of stalked eyes.
Atelocerata is a proposed clade of arthropods that includes Hexapoda and Myriapoda, but excludes Crustacea and Chelicerata. The name is currently used interchangeably with Tracheata. or Uniramia sensu stricto. It is an extensive division of arthropods comprising all those that breathe by tracheae, as distinguished from Crustacea, which breathe by means of gills.
Speleonectidae is a family of remipedes in the order Nectiopoda. There are at least two genera and about seven described species in Speleonectidae.
Godzilliidae is a family of remipedes in the order Nectiopoda. There are at least two genera and four described species in Godzilliidae.
The subphylum Mandibulata constitutes one of the major subdivisions of the phylum Arthropoda, alongside Chelicerata. Mandibulates include the crustaceans, myriapods, and all true insects. The name "Mandibulata" refers to the mandibles, a modified pair of limbs used in food processing, the presence of which are characteristic of most members of the group.
The Myriochelata or Paradoxopoda, is a proposed grouping of arthropods comprising the Myriapoda and Chelicerata. If this proposition holds true, the Myriochelata are the sister clade to the Pancrustacea, comprising classic crustaceans and hexapods.
Arthropods are invertebrates in the phylum Arthropoda. They possess an exoskeleton with a cuticle made of chitin, often mineralised with calcium carbonate, a body with differentiated (metameric) segments, and paired jointed appendages. In order to keep growing, they must go through stages of moulting, a process by which they shed their exoskeleton to reveal a new one. They form an extremely diverse group of up to ten million species.
Xenocarida is a proposed clade inside the subphylum Crustacea that comprises two classes that were discovered in the 20th century: Remipedia and Cephalocarida. Both groups are marine hermaphrodites. The clade was recovered as the sister groups to Hexapoda.
The subphylum Hexapoda or hexapods comprises the largest clade of arthropods and includes most of the extant arthropod species. It includes the crown group class Insecta, as well as the much smaller clade Entognatha, which includes three classes of wingless arthropods that were once considered insects: Collembola (springtails), Protura (coneheads) and Diplura. The insects and springtails are very abundant and are some of the most important pollinators, basal consumers, scavengers/detritivores and micropredators in terrestrial environments.
Tactopoda or Arthropodoidea is a proposed clade of ecdysozoan animals that includes the phyla Tardigrada and Euarthropoda, supported by various morphological observations. The cladogram below shows the relationships implied by this hypothesis.
Crustaceans are invertebrate animals that constitute one group of arthropods that are a part of the subphylum Crustacea, a large, diverse group of mainly aquatic arthropods including decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, opossum shrimps, amphipods and mantis shrimp. The crustacean group can be treated as a subphylum under the clade Mandibulata. It is now well accepted that the hexapods emerged deep in the Crustacean group, with the completed pan-group referred to as Pancrustacea. The three classes Cephalocarida, Branchiopoda and Remipedia are more closely related to the hexapods than they are to any of the other crustaceans.
The invertebrate mitochondrial code is a genetic code used by the mitochondrial genome of invertebrates. Mitochondria contain their own DNA and reproduce independently from their host cell. Variation in translation of the mitochondrial genetic code occurs when DNA codons result in non-standard amino acids has been identified in invertebrates, most notably arthropods. This variation has been helpful as a tool to improve upon the phylogenetic tree of invertebrates, like flatworms.
The clade Multicrustacea constitutes the largest superclass of crustaceans, containing approximately four-fifths of all described non-hexapod crustacean species, including crabs, lobsters, crayfish, shrimp, krill, prawns, woodlice, barnacles, copepods, amphipods, mantis shrimp and others. The largest branch of multicrustacea is the class Malacostraca.
The Hexanauplia is a clade proposed by Oakley et al. (2013) that constitutes a class of crustaceans, comprising the Copepoda and Thecostraca. A number of recent phylogenomic studies have not found support for this clade, with some supporting the alternative clade of Communostraca comprising Thecostraca and Malacostraca.
Allotriocarida is a clade of Pancrustacea, containing Hexapoda. It also contains three non-hexapod classes: Remipedia, Cephalocarida, and Branchiopoda. Newer studies also relocate copepoda, which traditionally has belonged to the multicrustaceans, to the Allotriocarida.