The halkieriids are a group of fossil organisms from the Lower to Middle Cambrian. Their eponymous genus is Halkieria , which has been found on almost every continent in Lower to Mid Cambrian deposits, forming a large component of the small shelly fossil assemblages. The best known species is Halkieria evangelista, from the North Greenland Sirius Passet Lagerstätte, in which complete specimens were collected on an expedition in 1989. The fossils were described by Simon Conway Morris and John Peel in a short paper in 1990 in the journal Nature. Later a more thorough description was undertaken in 1995 in the journal Philosophical Transactions of the Royal Society of London and wider evolutionary implications were posed.
The group is sometimes equated to Sachitida, although as originally envisaged, this group includes the wiwaxiids and is thus equivalent to the Halwaxiida.
History of discovery
Armor plates called "sclerites" had long been known as elements of the small shelly fossils, and detailed analysis showed that some of these belonged to the same animal and how they fitted together. The first articulated specimens of Halkieria evangelista, with all their hard parts together, were collected in 1989 from the Sirius Passet lagerstätte in Greenland, and were described in 1990 by Simon Conway Morris and John S. Peel. H. evangelista is used as a model for identifying and reconstructing as halkieriids other similar shells and sclerites; its epithet evangelista reflects its power to explain the Lower Cambrian fossil record.
Description of the fossils
Features shared by Halkieria and Australohalkieria
Only armor-like sclerites of Australohalkieria have been found, and much of the analysis assumes that these animals were similar to Halkieria. However the sclerites are so similar that this assumption looks fairly safe. In both genera the sclerites are of the type called "coelosclerites", which have a mineralized shell around a space originally filled with organic tissue, and which show no evidence of growth by adding material round the outside. Both genera also have sclerites of three different shapes: "palmate", flat and shaped rather like a maple leaf, which are generally the smallest; "cultrate", flat but shaped like knife blades; and "siculate", which are about the same size as the cultrates but are spine-shaped and like rather squashed cylinders. In both Halkieria and Australohalkieria the palmate and cultrate sclerites have prominent ribs, and are fairly flat except for 90° bends at the bases, which indicate that they fitted snugly against the animals' bodies. The siculates mostly lack ribs and appear to have projected away from the body at angles between about 45° and 90°.
Halkieria evangelista
The animals looked like slugs in chain mail - to long, bilaterally symmetric, flattened from top to bottom and unarmored on the bottom. Very near each end there is a shell plate with prominent growth lines rather like the growth rings of trees. The rest of the upper surface was covered with about 2,000 sclerites that overlapped each other like tiles and formed three zones with sclerites of different shapes: "palmates", shaped rather like maple leaves, ran along the center of the back between the shell plates; blade-shaped "cultrates" lay on either side of the palmates and pointing towards the middle of the upper surface; and slim, sickle-shaped "siculates" covered the outer edges. The sclerites bore a wide central cavity, and (at least in some specimens) finer lateral canals. As the animals grew, the shell plates grew by adding material to the outer edges. Individual sclerites stayed the same size; since the cultrate sclerites form a pattern that is constant in all fairly complete specimens, the old ones that were too small may have been shed and replaced by larger ones as the animals grew. The sclerites seem to have grown by basal secretion. There are traces of thin ribs between the sclerites and the skin.
The shellplates and the sclerites were probably made of calcium carbonate originally; it has been suggested on the basis of how they were preserved that they may have been wholly organic, but this is less likely since fossils of non-calcified organisms are usually thin films while Halkeieria fossils are three-dimensional like those of trilobites and hyoliths - in fact several specimens show curvature in the horizontal plane, which suggests that the muscles associated with the sclerites were still present at the time of burial