Tuesday, December 6, 2016

Wormworld: Biological Transitions At The Precambrian-Cambrian Boundary

The earliest animals were worms and they had a profound impact on marine ecosystems.

The many theories and some new understanding on the always fascinating topic of early animal evolution has been summarized quite well in a paper by James Schiffbauer and colleagues.

Molecular divergence time estimates (e.g., Erwin et al., 2011;Peterson et al., 2008) suggest that the last common ancestor of all animals evolved in the Cryogenian (ca. 800 Ma; although see dos Reis et al., 2015, for caveats). The earliest interpreted stem-group animals, however, are the ca. 600 Ma Doushantuo embryo-like microfossils (Chen et al., 2014a; Yin et al., 2016), leaving a
200-m.y. interlude between the fossil and molecular records. This hiatus between the estimated origin of Metazoa and their first appearance in the fossil record highlights the growing realization that the earliest stages of animal diversification were neither truly Cambrian nor explosive—with the phylogenetic origin of animals temporally removed from their morphological and ecological diversification by a long fuse (e.g., Conway Morris, 2000; Xiao, 2014). 

In this case, the significant lag between the establishment of the developmental toolkits necessary for the origin of novelty and their later implementation and ecological success can perhaps be attributed to the uniqueness of newly developing animal ecosystems. Between the ignition of the fuse and the subsequent evolutionary boom, three major eco-environmental feedbacks (see Erwin et al., 2011) arose that helped to pave the way for the Cambrian Explosion: (1) linkages between the pelagic and benthic ecosystems; (2) expansion of ecosystem engineering; and (3) metazoan macropredation. These feedbacks are explored herein in the context of the terminal Ediacaran fossil
record of vermiform organisms. This “wormworld” biota— comprised of various tubicolous body fossils (Figs. 2A–2C), such as the cloudinids, and increasingly complex vermiform ichnofossils (Figs. 2D–2F)—critically occupied a fundamental phase shift from competition- to predation-governed marine benthic ecosystems.

What was the big change in macroscopic life habits from the Precambrian to Cambrian times? Macroscopic multicellular life of the Ediacaran was dominated by benthic sessile forms. Early Cambrian animals were mobile creatures engaged in predation, burrowing, grazing and reef building. These activities resulted in an ecosystem engineering of sorts. For example; a) grazing and burrowing activity churned up sediment and oxygenated it. b) the evolution of guts in bilaterians transferred nutrients from sea water to the sediment in the form of fecal pellets.  These life modes created new ecologic niches and opened up new potential evolutionary pathways.

..And what killed out the classic Ediacaran biota. Was is environmental changes or ecologic competition from early animals?

It is important to note that the suggested mass extinction of the Ediacara biota in the context of our wormworld model is an ecologically driven event rather than an environmentally driven cataclysm akin to more recent (Phanerozoic) mass extinctions, and thus may have been comparatively protracted—as evidenced by Ediacara holdovers in the early Cambrian (Conway Morris, 1993; Hagadorn et al., 2000; Jensen et al., 1998). Nonetheless, whereas the static synecology and comparatively passive feeding modes of the classic Ediacarans had once emplaced a boundary on evolutionary possibility, the successful expansion of innovative traits of herbivory and carnivory, and their causal ties to infaunalization, reef-building, and biomineralization, permitted a new scaling of this bounding “right wall” (sensu Knoll and Bambach, 2000) as realized by the organisms of the wormworld fauna. Over time, the evolutionary breakthroughs conveyed by these neoteric organisms, including novel strategies, behaviors, and physiologies, increased the heterogeneity of benthic ecosystems, allowed for enhanced exploitation of resources, and established insurmountable increases in ecospace that ultimately signaled the curtain call for the Ediacara-type guilds.

The question of extinction of Ediacaran biota though may be more open ended than that suggested in this paper. E F Smith and colleagues in a recent issue of Geology analyze carbon isotope signatures of a carbonate succession spanning the Precambrian-Cambrian boundary. They find the carbonate sediment have pronounced negative carbon isotope values signalling a collapse or significant decrease in primary productivity in the oceans. 

What is the link between ecosystem collapse and negative carbon isotope excursion in carbonate sediment? Organic tissue preferentially incorporates C12, the lighter isotope of carbon. That means in thriving ecosystems, life is using up C12 from sea water and less of it makes its way into growing CaCO3 crystals forming carbonate sediment on the sea floor. When ecosystems collapse due to a myriad of reasons resulting in mass extinction, there is more C12 available to get incorporated into carbonate sediment. This increase in the lighter isotope C12 is a negative excursion of del13C, the ratio of C13 to C12.

Additional environmental disturbances may also contribute C12 to sea water. Warming of ocean water may lead to thawing of gas hydrates trapped below the sea bed. Methane released from hydrates is isotopically light and may break down and contribute C12 that eventually makes its way into carbonate. On land, a collapse of vegetation may release pulses of lighter carbon to the sea. Such a scenario would be realized in post Silurian times after the evolution of land vegetation.  In short, environmental catastrophe is linked to disturbances of the carbon cycle, and many sources may provide C12 to marine carbonate being formed at that time.

Anyways, what that means is that the decline in Ediacaran biota may have been due to both an environmental calamity as well as by longer term persistent competition by early animal activity.

And here is an infographic that summarizes the significant geological, ecological and biological events spanning the Precambrian-Cambrian transition

Source: Schiffbauer et al 2016

Open Access.

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