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Machaeridia is an extinct group of armoured, segmented
annelid The annelids (), also known as the segmented worms, are animals that comprise the phylum Annelida (; ). The phylum contains over 22,000 extant species, including ragworms, earthworms, and leeches. The species exist in and have adapted to vario ...
worms, known from the Early
Ordovician The Ordovician ( ) is a geologic period and System (geology), system, the second of six periods of the Paleozoic Era (geology), Era, and the second of twelve periods of the Phanerozoic Eon (geology), Eon. The Ordovician spans 41.6 million years f ...
(Late Tremadoc) to
Carboniferous The Carboniferous ( ) is a Geologic time scale, geologic period and System (stratigraphy), system of the Paleozoic era (geology), era that spans 60 million years, from the end of the Devonian Period Ma (million years ago) to the beginning of the ...
. It consists of three distinct families: the plumulitids, turrilepadids and lepidocoleids.


Fossils

Only the calcitic sclerites ("armour plates") of these worms tend to be preserved in the fossil record. These are tiny, and usually found disarticulated: articulated specimens reach about a centimeter in length, and are incredibly rare – hence the limited degree of study since their description in 1857. The machaeridians are characterized by having serialized rows of calcitic shell plates. The dorsal
sclerite A sclerite (Greek language, Greek , ', meaning "hardness, hard") is a hardened body part. In various branches of biology the term is applied to various structures, but not as a rule to vertebrate anatomical features such as bones and teeth. Instea ...
s were convex and almost isometric; lateral sclerites were flatter and longer. The plates comprised two calcite layers: the outer layer is thin and formed by lamellar deposition, whereas new elements were added to the thicker inner layer as it grew. Scales are ridged with growth lines, implying that they grew episodically. A few taxa experimented with different approaches to scale formation; some were only very weakly calcified and may have mainly been organic in nature. They were never moulted, and each scale could be moved with an attached muscle. The front two segments of the machaeridians were commonly different from the rest, bearing fewer spiny projections. The plumulitids are flattened from above and looks much like the coat of mail armour of chitons. The two other families are laterally compressed and some lepidocoleids formed a dorsal hinge, which make these machaeridians look like a string of bivalves.


Ecology

Machaeridians are often found in association with
stylophoran The stylophorans are an extinct, possibly polyphyletic group allied to the Paleozoic Era echinoderms, comprising the prehistoric cornutes and mitrates. It is synonymous with the subphylum Calcichordata. Their unusual appearances have led to a va ...
s - the
cornute Cornuta is an extinct order of echinoderms. Along with the mitrate Mitrates are an extinct group of stem group echinoderms, which may be closely related to the hemichordates. Along with the cornutes, they form one half of the Stylophora. ...
s and
mitrate Mitrates are an extinct group of stem group echinoderms, which may be closely related to the hemichordates. Along with the cornutes, they form one half of the Stylophora. Morphology The organisms were a few millimetres long. Like the echinod ...
s. This suggests that they possessed a similar ecology. They probably fed on organic detritus, perhaps even the
faeces Feces (also known as faeces American and British English spelling differences#ae and oe, or fæces; : faex) are the solid or semi-solid remains of food that was not digested in the small intestine, and has been broken down by bacteria in the ...
of the accompanying stylophorans. Their scales almost certainly performed a defensive role. The organisms would have had limited ability to flex to the right and left (in the
sagittal plane The sagittal plane (; also known as the longitudinal plane) is an anatomical plane that divides the body into right and left sections. It is perpendicular to the transverse and coronal planes. The plane may be in the center of the body and divi ...
), but would have been able to roll up. While most possessed bilateral symmetry, the scales on the right and left side of ''Turrilepas wrightiana'' are different in shape and form. The Plumulitid machaeridians would have moved across the surface of the sea floor using parapodia, whereas the fully armoured Turrelepids and Lepidocoelids burrowed in a peristaltic fashion reminiscent of their evolutionary cousins, the earthworms. This burrowing role has subjected them to the same evolutionary pressures which affect burrowing bivalves;
convergent evolution Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last comm ...
as a result of their shared function probably contributed to early suggestions that the machaeridians should be classified with the molluscs.


Taxonomic affinity

Historically the group has been assigned to the echinoderms, barnacles, annelids and mollusks. Relationships to other Cambrian forms (such as the Halkieriids) have been proposed and discounted. In 2008, the discovery of a fossil preserving soft tissue (including chaetae and parapodia) established an annelid affinity. Machaeridians represent the only instance of this group developing calcitic armour (notwithstanding certain polychaetes that integrate calcite into their chaetae). The exact position with annelids remains unresolved, though some characters indicate a relationship to Aphroditacean annelids (Vinther et al. 2008). In an accompanying commentary, Jean-Bernard Caron suggested that machaeridians must be a stem group based on number of specialised features. However, one cannot assess crown group/stem group affinities based on autapomorphies, but on shared morphological traits or the lack thereof. He also suggested that machaeridians might be polyphyletic, but machaerdians are a well defined group with a number of shared characters and morphological gradations among all three families. A study in 2019 recognized machaeridians as phyllodocids based on their jaws.


Articulated specimens

Articulated machaeridians are known from: ... and possibly elsewhere


References

{{Taxonbar, from=Q6723361 Annelids Controversial taxa Prehistoric annelids
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