Ordovician echinoderms

Exceptionally preserved fossils yield crucial information about the evolution of Life on Earth. The Fezouata Biota from the Lower Ordovician of Morocco is a Konservat-Lagerstätte of major importance, and it is today considered as an 'Ordovician Burgess Shale.' This biota was discovered only some 15 years ago, but geological studies of the area date back to the beginning of the 20th century. Pioneering geological investigations lead to the discovery of Ordovician strata in the Anti-Atlas (1929) and ultimately resulted in their formal subdivision into four main strat-igraphic units (1942). In the Agdz area, the presence of fossiliferous Tremadocian (Lower Ordovician) strata was suspected as early as 1939, but only definitively confirmed in 1955. In the 1960s–1990s, Jacques Destombes provided the first detailed biostratigraphic framework for the Lower Ordovician of the Anti-Atlas, and collected thousands of fossils that were subsequently described in a series of monographs. In the early 2000s, exceptionally preserved fossils were discovered in the Fezouata Shale (Tremadocian–late Floian) in the central Anti-Atlas by Mohamed 'Ou Saïd' Ben Moula. This biota, now known as the Fezouata Biota, is of utmost importance, for it demonstrates the extent in the fossil record of non-biomineralising animals typical of the 'Cambrian Explosion' into the Ordovician, during the 'Great Ordovician Biodiversification.' Although most components are still in need of formal descriptions, a fairly good picture of the composition and organisation of this biota, and how it contributes to our understanding of the early evolution of metazoan communities, can now be depicted. Moreover, recent studies have substantially clarified the biostratigraphical and palaeoenvironmental context of the Fezouata Shale, and are now being followed up by detailed investigations of the taphonomy, geochemistry and micropalaeontology of this unique Konservat-Lagerstätte. These efforts will soon greatly benefit from the recovery of fresh, unweathered samples from drill cores.

ABSTRACT Pelmatozoans diversified primarily during the Middle and Late Ordovician Period, with Early Ordovician records being much more limited, resulting in many gaps in our knowledge of the early stages of their diversification. Dissociated pelmatozoan ossicles have been found abundantly in one section in the Tonggao Formation (Tetragraptus approximatus Biozone, Floian, Early Ordovician). Most of the ossicles are thecal plates and stem ossicles from hemicosmitoid and glyptocystitoid cystoids. Thecal plates of ‘Cheirocrinus’ sp., Polycosmites sp., and other plates of uncertain affinity are described. A different ossicle type, Pentagonopentagonalis (col.), may represent crinoid remains; this would be one of the earliest occurrences of the class. The thecal ossicles and columnals are all considered, as both sets of data are desirable to determine the most complete estimate of generic diversity. The echinoderm ossicles may have been transported in from shallower water palaeoenvironments and clusters of ossicles may represent coprolites or regurgitates. Estimates of Early Ordovician palaeogeography that place this site at 30°S or near the palaeoequator are supported by the physiological requirements of the primitive echinoderms described herein. Copyright © 2014 John Wiley & Sons, Ltd.

The comment by Smith et al. rejects a previous interpretation of Middle Ordovician fossils as holothurians. This rejection is based on the use of inappropriate techniques to study the fossils, and insufficient consideration of their taphonomy. The fossils can be shown not to be sponges (the suggested alternative interpretation), and despite the taphonomic limitations, do show echinoderm characters such as the pentagonal oral ring and ambulacral structures.