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Halwaxiida

Halwaxiida is a proposed clade equivalent to the older orders Sachitida He 1980 and Thambetolepidea Jell 1981, loosely uniting scale-bearing Cambrian animals, which may lie in the stem group to molluscs or lophotrochozoa. Some palaeontologists question the validity of the Halwaxiida clade.

Definition and members
The name "halwaxiid" was formed by combining the names of two members of the proposed group, Halkieria and Wiwaxia. The group was defined as a set of Early to Mid Cambrian animals that had: a "chain mail" coat of three concentric bands of small armor plates that are called sclerites; in some cases, a small cap-like shell at the front end and in some cases both ends. and that their similarities are not biologically equivalent but evolved separately. In fact the originators of the term "halwaxiid" prefer an evolutionary "family tree" in which the halwaxiids are not monophyletic. Other fossil organisms became involved in the debate as there were plausible arguments for considering them closely related to the halkieriids or Wiwaxia or both. Orthrozanclus, first described in 2007, looked an intermediate between the halkieriids and Wiwaxia as it had a shell like that of halkieriids, and unmineralized sclerites and long spines like those of Wiwaxia – in fact the article which first described Orthrozanclus introduced the term "halwaxiid". New finds of Odontogriphus, reported in 2006, put this animal into play as well – despite its lack of sclerites or shells, its feeding apparatus looks very like Wiwaxia's. The siphogonuchitids, a very Early Cambrian group known only from isolated sclerites among the small shelly fossils, also appear in analyses as their sclerites suggest this group may have been close to the ancestors of halkieriids.The original, possibly incomplete, list includes Australohalkieria, Drepanochites, Eohalobia, Halkieria, Lomasulcachites, Ninella, Ocruranus, Oikozetetes, Orthrozanclus, Sinosachites, Siphogonuchites, Thambetolepis, and Wiwaxia — {{cite journal == Ecology ==
Ecology
Halwaxiid sclerites were not able to grow once they had been formed, but must rather have been either moulted or resorbed to make way for new, larger sclerites; the smallest Wiwaxia specimens, 3.4 mm in length, appear to bear the same number of sclerites (minus the spines) as adults 5 cm long. The new sclerites cannot simply have poked between existing sclerites; rather they must have been shed or resorbed, either one at a time or all at once; presumably the latter, as the close-fitting nature of the armour does not seem compatible with the easy loss of individual sclerites. ==Suggested phylogenies==
Suggested phylogenies
Since 1995 several phylogenies, or evolutionary "family trees", have been suggested for these organisms. The position of Wiwaxia is highlighted, since the position of this organism has been central to the debate since 1990. propose Halkieria as a stem brachiopod, and Wiwaxia as a stem mollusc. ==Decline==
Decline
Wiwaxia and Halkieria did not thrive for a long period of time and were extinct before the Cambrian was over. A possible reason for this is that during the Cambrian substrate revolution, the microbial mat's disappearance triggered an alternating pattern in the fauna that were present for the Halwaxiids to eat. ==Wiwaxia and Odontogriphus==
Wiwaxia and Odontogriphus
feeding apparatus (1985): When he briefly described the first articulated specimens of Halkieria'' in 1990, Conway Morris wrote of "the halkieriid-wiwaxiid body plan" and that the halkieriids might be close relatives of molluscs. {{Annotated image | float=right | caption=Halkieriid sclerite structure Recent finds indicate that Orthrozanclus is likely a halkieriid, but has little affinity to Wiwaxia. ==Halkieriids and siphogonuchitids==
Halkieriids and siphogonuchitids
Most of the debate has been about halkieriids' relationship to molluscs, annelids and brachiopods. However a paper published in 2008 argued that halkieriids were closely related to chancelloriids, which have been generally regarded as sponges. Relationship to Molluscs, Annelids and Brachiopods Halkieriid sclerites were known for a long time as elements of the small shelly fossils. The first articulated specimen, with all their hard parts together, were collected in 1989 and were described briefly in 1990 by Conway Morris and Peel (1990). Vinther and Nielsen (2005) proposed instead that Halkieria was a crown group mollusc, in other words more similar to modern molluscs that to annelids, brachiopods or any intermediate groups. They argued that: Halkieria's sclerites resembled those of the modern solenogaster aplacophoran shell-less molluscs (see ), of some modern polyplacophoran molluscs, which have several shell plates, and of the Ordovician polyplacophoran Echinochiton; Halkieria's shells are more similar to the shells of conchiferan molluscs, since shells of both of these groups show no trace of the canals and pores seen in polyplacophoran shell plates; the bristles of brachiopods and annelids are similar to each other but not to Halkieria's sclerites. However Conway Morris (2006) criticized Vinther and Nielsen's classification of Halkieria as a crown group mollusc, on the grounds that the growth of the spicules in the aplacophorans and polyplacophorans is not similar to the method of growth deduced for the complex halkieriid sclerites; in particular, he said, the hollow spines of various molluscs are not at all like the halkieriid sclerites with their complex internal channels. Conway Morris repeated his earlier conclusion that halkieriids were close to the ancestors of both molluscs and brachiopods. In their description of the newly discovered Orthrozanclus (2007), which has similarities to both Wiwaxia and the halkieriids, Conway Morris and Caron also took account of the siphogonuchitids, a group known only from isolated mineralized sclerites that resemble those of the halkieriids. They proposed the two "family trees" described above: • The halkieriids formed part of a "sister" group to the molluscs. This hypothesis implies that the halwaxiids were not monophyletic, since the siphogonuchitids appear between Wiwaxia and the halkieriids. Nevertheless, Conway Morris and Caron found that this phylogeny fitted the available data better, although it was not robust. • The siphogonuchitids plus the halwaxiids formed a "sister" group to the annelids and brachiopods, while the molluscs were "aunts" of all of these. Porter (2008) found that the sclerites of halkieriids and chancelloriids resemble each other at all levels: both have an internal "pulp cavity" and a thin external organic layer; the walls are made of the same material, aragonite; the arrangement of the aragonite fibers is in each is the same, running mainly from base to tip but with each being closer to the surface at the end nearest the tip. It is extremely improbable that totally unrelated organisms could have developed such similar sclerites independently, but the huge difference in the structures of their bodies makes it hard to see how they could be closely related. This dilemma may be resolved in various ways: but shows no evidence of sclerites. ==Notes and references==
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