Expression of Hairpin-Enriched Mitochondrial DNA in Two Hairworm Species (Nematomorpha).

Nematomorpha (hairworms) is a phylum of parasitic ecdysozoans, best known for infecting arthropods and guiding their hosts toward water, where the parasite can complete its life cycle. Over 350 species of nematomorphs have been described, yet molecular data for the group remain scarce. The few available mitochondrial genomes of nematomorphs are enriched with long inverted repeats, which are embedded in the coding sequences of their genes-a remarkably unusual feature exclusive to this phylum. Here, we obtain and annotate the repeats in the mitochondrial genome of another nematomorph species-Parachordodes pustulosus. Using genomic and transcriptomic libraries, we investigate the impact of inverted repeats on the read coverage of the mitochondrial genome. Pronounced drops in the read coverage coincide with regions containing long inverted repeats, denoting the 'blind spots' of short-fragment sequencing libraries. Phylogenetic inference with the novel data reveals multiple disagreements between the traditional system of Nematomorpha and molecular data, rendering several genera paraphyletic, including Parachordodes.

Cambrian radiation speciation events driven by sea level and redoxcline changes on the Siberian Craton.

The evolutionary processes of speciation during the Cambrian radiation and their potential extrinsic drivers, such as episodic oceanic oxygenation events, remain unconfirmed. High-resolution temporal and spatial distribution of reef-associated archaeocyath sponge species on the Siberian Craton during the early Cambrian [ca. 528 to 510 million years ago] shows that speciation was driven by increased endemism particularly ca. 521 million years (59.7% endemic species) and 514.5 million years (65.25% endemic species) ago. These mark rapid speciation events after dispersal of ancestors from the Aldan-Lena center of origin to other regions. These speciation events coincided with major sea-level lowstands, which we hypothesize were intervals when relative deepening of the shallow redoxcline permitted extensive oxygenation of shallow waters over the entire craton. This provided oxic corridors for dispersal and allowed the formation of new founder communities. Thus, shallow marine oxygen expansion driven by sea-level oscillations provides an evolutionary driver for sucessive speciation events during the Cambrian radiation.

Increases in reef size, habitat and metacommunity complexity associated with Cambrian radiation oxygenation pulses.

Oxygenation during the Cambrian Radiation progressed via a series of short-lived pulses. However, the metazoan biotic response to this episodic oxygenation has not been quantified, nor have the causal evolutionary processes been constrained. Here we present ecological analyses of Cambrian archaeocyath sponge reef communities on the Siberian Platform (525-514 Ma). During the oxic pulse at ~521-519 Ma, we quantify reef habitat expansion coupled to an increase in reef size and metacommunity complexity, from individual within-community reactions to their local environment, to ecologically complex synchronous community-wide response, accompanied by an increase in rates of origination. Subsequently, reef and archaeocyath body size are reduced in association with increased rates of extinction due to inferred expanded marine anoxia (~519-516.5 Ma). A later oxic pulse at ~515 Ma shows further reef habitat expansion, increased archaeocyath body size and diversity, but weaker community-wide environmental responses. These metrics confirm that oxygenation events created temporary pulses of evolutionary diversification and enhanced ecosystem complexity, potentially via the expansion of habitable space, and increased archaeocyath individual and reef longevity in turn leading to niche differentiation. Most notably, we show that progression towards increasing biodiversity and ecosystem complexity was episodic and discontinuous, rather than linear, during the Cambrian Radiation.

The oldest mineralized bryozoan? A possible palaeostomate in the lower Cambrian of Nevada, USA.

All skeletal marine invertebrate phyla appeared during the Cambrian explosion, except for Bryozoa with mineralized skeletons, which first appear in the Early Ordovician. However, the skeletal diversity of Early Ordovician bryozoans suggests a preceding interval of diversification. We report a possible earliest occurrence of palaeostomate bryozoans in limestones of the Cambrian Age 4 Harkless Formation, western United States. Following recent interpretations of the early Cambrian Protomelission as a soft-bodied bryozoan, our findings add to the evidence of early Cambrian roots for the Bryozoa. The Harkless fossils resemble some esthonioporate and cystoporate bryozoans, showing a radiating pattern of densely packed tubes of the same diameter and cross-sectional shape. Further, they show partitioning of new individuals from parent tubes through the formation of a separate wall, a characteristic of interzooecial budding in bryozoans. If confirmed as bryozoans, these fossils would push back the appearance of mineralized skeletons in this phylum by ~30 million years and impact interpretations of their evolution.

Dynamic and synchronous changes in metazoan body size during the Cambrian Explosion.

Many aspects of the drivers for, and evolutionary dynamics of, the Cambrian Explosion are poorly understood. Here we quantify high-resolution changes in species body size in major metazoan groups on the Siberian Platform during the early Cambrian (ca. 540-510 Million years ago (Ma)). Archaeocyath sponges, hyolith lophophorates, and helcionelloid mollusc species show dynamic and synchronous trends over million-year timescales, with peaks in body size during the latest Tommotian/early Atbadanian and late Atdabanian/early Botoman, and notably small body sizes in the middle Atdabanian and after the Sinsk anoxic extinction event, starting ca. 513 Ma. These intervals of body size changes are also mirrored in individual species and correlate positively with increased rates of origination and broadly with total species diversity. Calcitic brachiopods (rhynchonelliformeans), however, show a general increase in body size following the increase in species diversity through this interval: phosphatic brachiopods (linguliformeans) show a body size decrease that negatively correlates with diversity. Both brachiopod groups show a rapid recovery at the Sinsk Event. The synchronous changes in these metrics in archaeocyath, hyoliths and helcionelloids suggest the operation of external drivers through the early Cambrian, such as episodic changes in oxygenation or productivity. But the trends shown by brachiopods suggests a differing physiological response. Together, these dynamics created both the distinct evolutionary record of metazoan groups during the Cambrian Explosion and determined the nature of its termination.

Possible links between extreme oxygen perturbations and the Cambrian radiation of animals.

The role of oxygen as a driver for early animal evolution is widely debated. During the Cambrian explosion, episodic radiations of major animal phyla occurred coincident with repeated carbon isotope fluctuations. However, the driver of these isotope fluctuations and potential links to environmental oxygenation are unclear. Here, we report high-resolution carbon and sulphur isotope data for marine carbonates from the southeastern Siberian Platform that document the canonical explosive phase of the Cambrian radiation from ~524 to ~514 Myr ago. These analyses demonstrate a strong positive covariation between carbonate δ13C and carbonate-associated sulphate δ34S through five isotope cycles. Biogeochemical modelling suggests that this isotopic coupling reflects periodic oscillations in atmospheric O2 and the extent of shallow ocean oxygenation. Episodic maxima in the biodiversity of animal phyla directly coincided with these extreme oxygen perturbations. Conversely, the subsequent Botoman-Toyonian animal extinction events (~514 to ~512 Myr ago) coincided with decoupled isotope records that suggest a shrinking marine sulphate reservoir and expanded shallow marine anoxia. We suggest that fluctuations in oxygen availability in the shallow marine realm exerted a primary control on the timing and tempo of biodiversity radiations at a crucial phase in the early history of animal life.

The two phases of the Cambrian Explosion.

The dynamics of how metazoan phyla appeared and evolved - known as the Cambrian Explosion - remains elusive. We present a quantitative analysis of the temporal distribution (based on occurrence data of fossil species sampled in each time interval) of lophotrochozoan skeletal species (n = 430) from the terminal Ediacaran to Cambrian Stage 5 (~545 - ~505 Million years ago (Ma)) of the Siberian Platform, Russia. We use morphological traits to distinguish between stem and crown groups. Possible skeletal stem group lophophorates, brachiopods, and molluscs (n = 354) appear in the terminal Ediacaran (~542 Ma) and diversify during the early Cambrian Terreneuvian and again in Stage 2, but were devastated during the early Cambrian Stage 4 Sinsk extinction event (~513 Ma) never to recover previous diversity. Inferred crown group brachiopod and mollusc species (n = 76) do not appear until the Fortunian, ~537 Ma, radiate in the early Cambrian Stage 3 (~522 Ma), and with minimal loss of diversity at the Sinsk Event, continued to diversify into the Ordovician. The Sinsk Event also removed other probable stem groups, such as archaeocyath sponges. Notably, this diversification starts before, and extends across the Ediacaran/Cambrian boundary and the Basal Cambrian Carbon Isotope Excursion (BACE) interval (~541 to ~540 Ma), ascribed to a possible global perturbation of the carbon cycle. We therefore propose two phases of the Cambrian Explosion separated by the Sinsk extinction event, the first dominated by stem groups of phyla from the late Ediacaran, ~542 Ma, to early Cambrian stage 4, ~513 Ma, and the second marked by radiating bilaterian crown group species of phyla from ~513 Ma and extending to the Ordovician Radiation.

First macrobiota biomineralization was environmentally triggered.

Why large and diverse skeletons first appeared ca 550 Ma is not well understood. Many Ediacaran skeletal biota show evidence of flexibility, and bear notably thin skeletal walls with simple, non-hierarchical microstructures of either aragonite or high-Mg calcite. We present evidence that the earliest skeletal macrobiota, found only in carbonate rocks, had close soft-bodied counterparts hosted in contemporary clastic rocks. This includes the calcareous discoidal fossil Suvorovella, similar to holdfasts of Ediacaran biota taxa previously known only as casts and moulds, as well as tubular and vase-shaped fossils. In sum, these probably represent taxa of diverse affinity including unicellular eukaryotes, total group cnidarians and problematica. Our findings support the assertion that the calcification was an independent and derived feature that appeared in diverse groups where an organic scaffold was the primitive character, which provided the framework for interactions between the extracellular matrix and mineral ions. We conclude that such skeletons may have been acquired with relative ease in the highly saturated, high alkalinity carbonate settings of the Ediacaran, where carbonate polymorph was further controlled by seawater chemistry. The trigger for Ediacaran biomineralization may have been either changing seawater Mg/Ca and/or increasing oxygen levels. By the Early Cambrian, however, biomineralization styles and the range of biominerals had significantly diversified, perhaps as an escalating defensive response to increasing predation pressure. Indeed skeletal hardparts had appeared in clastic settings by Cambrian Stage 1, suggesting independence from ambient seawater chemistry where genetic and molecular mechanisms controlled biomineralization and mineralogy had become evolutionarily constrained.