Pleistoannelida | |
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Examples of Pleistoannelida (clockwise from upper left corner): Sabellida, Echiura, Eunicida, Clitellata, Phyllodocida, Siboglinidae. | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
(unranked): | Spiralia |
Superphylum: | Lophotrochozoa |
Phylum: | Annelida |
Clade: | Pleistoannelida Struck 2011 [1] |
Clades [2] [3] [4] | |
Incertae sedis |
Pleistoannelida is a group of annelid worms that comprises the vast majority of the diversity in phylum Annelida. Discovered through phylogenetic analyses, it is the largest clade of annelids, comprised by the last common ancestor of the highly diverse sister groups Errantia and Sedentaria (Clitellata and related polychaetes) and all the descendants of that ancestor. [1] [4] Most groups in the Clade find their ancestors within the Cambrian explosion when Annelid diversity expanded dramatically. [5] The Pleistoannelida clade covers a variety of traits. However, the evolution of simple to complex eyes, developed papillae for burrowing, and for some specialized radioles for feeding can be seen universally across every species. [6] [5] New findings have discovered the range of Annelid diversity have led to uncertainty if groups with developed ancestral traits should remain within the clade. Furthermore, there's been a lack of recently discovered Annelid traits being used in the categorization of groups within the clade, leading to many hypothesis on how to do so and which should remain within the clade. [7] Currently three smaller clades that were originally a part of the groups Errantia and Sedentaria have been proven to fall outside while still being connected to the basal groups. [6]
Pleistoannelida is composed by two big clades: Errantia (eunicids, siboglinids and related polychaetes) and Sedentaria (spoon worms, tubeworms, clitellates and related polychaetes). There are also smaller groups of difficult placement, such as Myzostomida and Spintheridae, that belong to either one of them. [2] Closely related to Pleistoannelida is a grade of basal annelids: the Amphinomida/Sipunculida/ Lobatocerebrum clade, Chaetopterida and Palaeoannelida. [4] The clade Amphinomida was once originally included within Errantia, but has since been removed into its own taxa.
A possible, closer sister group to Pleistoannelida could be Dinophiliformia, a clade containing interstitial (meiofaunal) genera previously found in Orbiniida (Sedentaria). [8]
A 2019 analysis recovered the mesozoan group Orthonectida as the sister group to Pleistoannelida. [9] However, a more recent 2022 analysis found a monophyletic Orthonectida+Dicyemida clade localized outside of the annelids, between Gnathifera and Platyhelminthes. [10]
There has been research conducted to help determine the taxonomy of the clade Pleistoannelida. The research includes the morphological characteristics of the clade as well as the phylum Annelida as a whole. For example, there are studies of gene expression in annelids. It was supported that larval eyes of annelids are considered homologous to pigmented eyes of bilaterians. It is believed that annelid adult eyes evolved in a common ancestor of Pleistoannelida. [11] Besides the eyes, it has also been studied that commissures in the brain, glanglia, and nuchal organs were to have also evolved in the lineage of Pleistonannelida (Errantia and Sedentaria) by observing Magelonamirabilis. [12] At a cellular level, it was found that the highly conserved mitochondrial gene order could be only depicted to the clade Pleistoannelida. [13]
There has been much research done on the genetics of Pleistoannelida and Annelida as a whole to determine the phylogeny of the clade, because many taxa do not share ancestral characteristics. Genetic studies have led to the current phylogenetic tree, which separates the clade into the two sister groups of Errantia and Sedentaria. [14] Genetic data was also used in the placement of groups of Myzostomida, Nerilliade, and Aberranta within Pleistoannelida. These groups were previously thought to be anywhere from annelids to flatworms, but genetics studies have found strong evidence to place them solidly in Annelida, with the closest relations to groups in the clade Pleistoannelida. [15] [16] The most current research puts Myzostomida in Errantia, however the data is not strong enough to make this a certainty. [13] Research places Nerillidae closest to Eunicida (supported by certain morphology) and places Aberranta near Nerillidae or syllids and nereidids. [15]
Siboglinidae is a family of polychaete annelid worms whose members made up the former phyla Pogonophora and Vestimentifera. The family is composed of around 100 species of vermiform creatures which live in thin tubes buried in sediment (Pogonophora) or in tubes attached to hard substratum (Vestimentifera) at ocean depths ranging from 100 to 10,000 m. They can also be found in association with hydrothermal vents, methane seeps, sunken plant material, and whale carcasses.
Polychaeta is a paraphyletic class of generally marine annelid worms, commonly called bristle worms or polychaetes. Each body segment has a pair of fleshy protrusions called parapodia that bear many bristles, called chaetae, which are made of chitin. More than 10,000 species are described in this class. Common representatives include the lugworm and the sandworm or clam worm Alitta.
The Trochozoa are a proposed Lophotrochozoa clade that is a sister clade of Bryozoa and Platyzoa. The clade would include animals in five phyla: the Nemertea, the Annelida, the Mollusca, and the two Brachiozoan phyla, Brachiopoda and Phoronida. Both annelids and molluscs have been suggested as the sister group of Brachiozoa. It has also been proposed that nemerteans are actually a clade of annelids.
Orthonectida is a small phylum of poorly known parasites of marine invertebrates that are among the simplest of multi-cellular organisms. Members of this phylum are known as orthonectids.
Lophotrochozoa is a clade of protostome animals within the Spiralia. The taxon was established as a monophyletic group based on molecular evidence. The clade includes animals like annelids, molluscs, bryozoans, and brachiopods.
The lophophore is a characteristic feeding organ possessed by four major groups of animals: the Brachiopoda, Bryozoa, Hyolitha, and Phoronida, which collectively constitute the protostome group Lophophorata. All lophophores are found in aquatic organisms.
The Clitellata are a class of annelid worms, characterized by having a clitellum – the 'collar' that forms a reproductive cocoon during part of their life cycles. The clitellates comprise around 8,000 species. Unlike the class of Polychaeta, they do not have parapodia and their heads are less developed.
The Myzostomida or Myzostomatida are an order of small marine worms, which are parasitic on echinoderms, mostly crinoids. These highly unusual and diverse annelids were first discovered by Friedrich Sigismund Leuckart in 1827.
The Spiralia are a morphologically diverse clade of protostome animals, including within their number the molluscs, annelids, platyhelminths and other taxa. The term Spiralia is applied to those phyla that exhibit canonical spiral cleavage, a pattern of early development found in most members of the Lophotrochozoa.
Rhodine is a genus of capitellid segmented worms in the family Maldanidae.
Errantia is a diverse group of marine polychaete worms in the phylum Annelida. Traditionally a subclass of the paraphyletic class Polychaeta, it is currently regarded as a monophyletic group within the larger Pleistoannelida, composed of Errantia and Sedentaria. These worms are found worldwide in marine environments and brackish water.
The annelids, also known as the segmented worms, comprise a large phylum called Annelida. It contains over 22,000 extant species, including ragworms, earthworms, and leeches. The species exist in and have adapted to various ecologies – some in marine environments as distinct as tidal zones and hydrothermal vents, others in fresh water, and yet others in moist terrestrial environments.
The Platytrochozoa are a proposed basal clade of spiralian animals as the sister group of the Gnathifera. The Platytrochozoa were divided into the Rouphozoa and the Lophotrochozoa. A more recent study suggests that the mesozoans also belong to this group of animals, as sister of the Rouphozoa.
Flabelligeridae is a family of polychaete worms, known as bristle-cage worms, notable for their cephalic cage: long slender chaetae forming a fan-like arrangement surrounding the eversible head. Unlike many polychaetes, they also have large, pigmented, complex eyes.
Rhodininae is a subfamily of marine polychaete worms in the family Maldanidae.
Sedentaria is a diverse clade of annelid worms. It is traditionally treated as a subclass of the paraphyletic class Polychaeta, but it is also a monophyletic group uniting several polychaetes and the monophyletic class Clitellata. It is the sister group of Errantia.
Lumbriclymeninae is a subfamily of marine polychaete worms in the family Maldanidae.
Notoproctus is a genus of marine polychaete worms in the family Maldanidae. It is the only member of the subfamily Notoproctinae.
Orbiniida is an order of small polychaete worms in the phylum Annelida. It is the earliest diverging clade in Sedentaria. Along with Protodriliformia, this order is composed of meiofaunal marine worms formerly known as "archiannelids". These worms inhabit the marine interstitial ecosystem, the space between sand grains.
Protodriliformia is a clade of small marine polychaetes, comprised by the groups of meiofaunal interstitial worms Protodrilida and Polygordiidae, formerly considered "archiannelids". It is the most basal clade of Errantia.