Conditions that may have underpinned the ‘Cambrian Explosion’

Geologists of my generation leaned that the earliest signs of abundant and diverse animal life were displayed by an extraordinary assemblage of fossils in a mudstone exposure high on a ridge in the Rocky Mountains of British Columbia. The Burgess Shale lagerstätte, or ‘site of exceptional preservation’, was discovered by Charles Walcott in 1909. It contained exquisite remains, some showing signs of soft tissue, of a great range of animals, many having never before been seen. Though dated at 509 Ma (Middle Cambrian) it was regarded for much of the 20th century as the sign of a sudden burgeoning from which all subsequent life had evolved: the Cambrian Explosion. Walcott only scratched the surface of its riches, its true wonders only being excavated and analysed later by Harry Whittington and his protégé Simon Conway Morris of Cambridge University. Their results were summarised and promoted in one of the great books on palaeontology and evolutionary biology, Wonderful Life (1989) by Steven Jay Gould.

Harbingers of animal profusion first appear around 635 Ma in the Late Neoproterozoic as the Ediacaran Fauna, with the oldest precursors turning up around a billion years ago in the Torridonian Sandstone Formation of northern Scotland. The evolutionary links between them and the Cambrian Explosion are yet to be documented, as creatures of the Ediacaran remain elusive in the earliest Phanerozoic rocks. As regards the conditions that promoted the explosion of animal faunas, the Burgess Shale is a blank canvas, for its riches were not preserved in situ, but had drifted onto deep, stagnant ocean floor to be preserved in oxygen-poor muds that enabled their intricate preservation. The animals could not have lived and evolved without abundant oxygen: what that environment was is not recorded by Walcott’s famous stratigraphic site.

Artistic impression of the Chengjian Biota

China, it has emerged, offers a major clue from around 40 lagerstätten in Chengjian County, Yunnan. They are not only older (518 Ma) than the Burgess Shale but contain 27 percent more faunal diversity: 17 phylums and more than 250 species. Since the discovery of the Chengjian Biota in the first decade of the 21st century palaeontologists have, understandably, been preoccupied by describing its riches in hundreds of scientific papers. The nature of the ecosystem has remained as obscure as that of the Burgess Shale, largely due to the exposed host rocks (laminated siltstones and mudstones) having been weathered. They are superficially similar to the Burgess Shale. In March 2022, 10 scientists working at laboratories in China, Canada, Switzerland and the UK published the results of their painstaking sedimentological investigation of a core dilled through through the entire fossiliferous sequence (Salih, F. and 9 others 2022. The Chengjiang Biota inhabited a deltaic environment. Nature Communications, v. 13, article 1569; DOI: 10.1038/s41467-022-29246-z).

Reconstruction of the near-shore deltaic environment in which the Chengjian Biota lived and evolved. Several rock types and the sedimentary processes that probably formed them shown in ‘cores’ (Credit: Salih et al. Figure 3)

The unweathered core displays a variety of tiny sedimentary structures. These include cross laminations formed by migrating ripples, occasional fine sandstones that include signs of burrowing, graded bedding formed by minor turbidity currents, hummocks formed by back and forth water flow, ripples formed by flow in a single direction and small channels. Unlike the Burgess Shale, the fine-grained Chengjian sediments seem to have been deposited in environments that were far from stagnant and deep. They most closely resemble the offshore parts of the delta of a predominantly muddy river, subject to occasional floods and storms and characterised by large and rapid accumulation of mud and silt by dense sediment-loaded river water flowing down a gently sloping seabed into clearer seawater. That the sediment supply was full of nutrients and oxygen is reflected by small organisms living in burrows. The high-quality preservation of fossils in some layers can be attributed to sudden influxes of freshwater into their marine habitat during storms, so that they were killed in place. Such a near-shore environment, full of nutrients and oxygen but subjected to repeated geochemical and physical stresses, can explain adaptive radiation and evolution at a fast pace. Clearly, that is by no means a full explanation of the Cambrian Explosion, but offers sufficient insight for research to proceed fruitfully.

See also: Modern Animal Life Could Have Origins in a Shallow, Nutrient-Rich Delta, SciTechDaily, 23 March 2022.

Hallucigenia gets a head

The Middle Cambrian Burgess Shale of the Canadian Rockies is one of those celebrated sediments that show extraordinary preservation of soft-bodied and easily disarticulated organisms and rich assemblages of fossils. Being one of the earliest known of such lagerstätten, many of the denizens of the ecosystem in which the shale originated were at first regarded as members of hitherto undiscovered and now vanished phyla, the basal branches of the ‘tree of life’. Some certainly looked pretty odd, such as Opabina with a feeding apparatus looking similar to the extension nozzle of a vacuum cleaner; but that is clearly some kind of arthropod. Others turned out to be astonishingly large, once it was realised that parts of their broken bodies had previously been taken to be different organisms, an example being Anomalocaris. But perhaps the oddest, certainly to palaeontologists, was Hallucigenia. However, there are plenty of even more weird and wonderful living creatures, such as the sea pig, although modern creatures are more easily pigeonholed, taxonomically speaking.

Halucigenia as originally reconstruicted (i.e....
Hallucigenia as originally reconstructed; i.e. upside-down. (credit: Wikipedia)

The trouble with Hallucigenia was not so much its complexity – it was a fairly simple-looking beast – but that there were two choices as to which way up it lived; a feature that surprisingly led to a great deal of pondering that ended with the scientist who formally described it in 1977 making the wrong choice. That was eventually resolved fourteen years later, but the creature might also have inspired the Pushmi Pullyu in Hugh Lofting’s Dr Doolittle stories for children. Not that it resembled a unicorn-gazelle cross: far from it, for no-one could decide which its front was and which its backside, and even if it may have lain on its side. But Hallucigenia does demonstrate bilateral symmetry beautifully – it must have a front and back, and a top and bottom, even though which was which remained veiled in mystery – and so belongs to the dominant group of animals, imaginatively known as bilaterians.

The Burgess Shale lagerstätte seemingly was heaving with Hallucigenia so would-be taxonomists have had no shortage of specimens to ponder over in the 38 years since Simon Conway Morris made his dreadful mistake: of course, that was not of such enormity as Einstein’s ‘biggest blunder’ in the form of his cosmological constant, and Conway Morris quickly accepted his error when the beast was turned right-way-up in 1991. The problem is, exquisite as they are, Burgess Shale fossils are flattened and all that remains of mainly soft-bodied animals are delicate carbonaceous films, which need electron microscopy to unravel.

The latest reconstruction of Hallucigenia, by palaeontological illustrator Danielle Dufault (http://www.ddufault.com)
The latest reconstruction of Hallucigenia, by palaeontological illustrator Danielle Dufault

In 2015, Hallucigenia’s front end was definitely found and a great deal more besides by Canadian palaeontologists Martin Smith and Jean-Bernard Caron of the Royal Ontario Museum and the University of Toronto (Smith, M.R. & Caron J.-B. 2015. Hallucigenia’s head and the pharyngeal armature of early ecdysozoans. Nature, v. 523, p. 75-78). It has eyes, albeit rudimentary, and a throat, deep within which it has pointy teeth. Hallucigenia was a lobopod, whose living relatives lie within that large and diverse group the Ecdysozoa, which all have throat teeth and include the wondrous water bear (tardigrade) and the velvet- and penis worms (onychophores and priapulids, respectively) as well as lobsters, flies and woodlice. It may indeed have been close to the last common ancestor of all animals who moult their carapaces.