Cambrian Explosion #34: Phylum Priapulida

Named for their resemblance to human penises, priapulids (or “penis worms”) are marine scalidophoran worms that live on or in muddy seafloor sediment, with some species having a surprisingly high tolerance for oxygen-poor environments and toxic levels of hydrogen sulfide. Despite being a rather low-diversity phylum with only around 20 living species, they’re widespread and sometimes very numerous, with over 80 adult individuals per square meter (~10ft²) recorded in some locations.

The earliest definite modern-style priapulid in the fossil record comes from the late Carboniferous (~308 million years ago), but their ancestry was probably somewhere in the early Cambrian among the taxonomic mess of palaeoscoloecids and archaeopriapulids.

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Cambrian Explosion #33: Early Scalidophora

The earliest branches of the ecdysozoan evolutionary tree are made up of the scalidophorans – animals with spiny retractable proboscises, represented today by the worm-like priapulids and kinorhynchs, and (sometimes*) the weird little loriciferans.

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Cambrian Explosion #32: Rise of the Arthropods

The world of the Cambrian Period was a strange combination of both familiar and alien. The land would have seemed rather barren, populated mainly by microbes and algae, yet the oceans teemed with creatures already identifiable as sponges, comb jellies, jellyfish, acorn worms, vertebrates, echinoderms, arrow worms, annelids, molluscs, and brachiopods – small and primitive-looking in some cases but still recognizable enough.

But at the same time there were “weird wonders” everywhere, things much harder to identify, with shapes so bizarre that their initial discovery was met with laughter.

Animal life was exploring so many different possibilities for body plans and ecologies, and one lineage in particular dominated this explosion of evolutionary experimentation: the arthropods.

Arthropods are represented today by the chelicerates (sea spiders, horseshoe crabs, and arachnids), myriapods (millipedes and centipedes), crustaceans, and insects, and together these groups make up over 80% of all known living animal species and are vital parts of almost every ecosystem on the planet.

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Nesonektris

Nesonektris aldridgei here was one of the bizarre vetulicolians, a group of Cambrian animals that lived between about 520 and 505 million years ago.

Known from the Emu Bay Shale fossil deposits in Kangaroo Island, South Australia (~514 million years ago), Nesonektris was one of the larger known vetulicolians, growing to at least 17cm long (~6.5″). Like most of its relatives it had a large streamlined forebody with a mouth opening at the front, and no obvious appendages or sensory structures. A groove down each side may have housed gill openings, and a segmented flexible tail provided propulsion for swimming.

Very little is known about the ecology of these animals. They were clearly adapted for active swimming in the water column, and may have filter-fed on plankton – but some other vetulicolians have been found preserved with their guts full of seafloor sediment, suggesting some sort of detritivorous lifestyle instead.

Their evolutionary relationships are also still uncertain, but preservation of what appears to be a notochord in Nesonektris suggests that vetulicolians may have been part of the chordate lineage, possibly close relatives of tunicates.

Cambrian Explosion Month #31: Phylum Brachiopoda

While modern brachiopods superficially resemble clams, they’re not actually very closely related to each other. Clams are bivalve molluscs, related to snails and squid, while brachiopods are lophophorates related to bryozoans and horseshoe worms.

Their two shell valves are also arranged very differently – while bivalve shells originate from the left and right sides of their bodies, brachiopods grow theirs on the top and bottom.

They first appear in the fossil record in the early Cambrian, about 530 million years ago, but they may have actually diverged from a tommotiid-like ancestor as far back as the late Ediacaran. Only around 300 species survive today, but during the Paleozoic they were some of the most abundant filter-feeding and reef-building animals with tens of thousands of fossil species known. Different species tended to have strict habitat and temperature preferences, and so their fossils are also useful indicators of how ancient climates changed over time.

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Cambrian Explosion Month #30: Phylum(?) Hyolitha

Hyoliths were a group of small shelled animals that first appeared in the fossil record just after the start of the Cambrian, about 536 million years ago. They had conical calcareous shells with a lid-like operculum, and some species also featured long curling spines that made them look like ice-cream cones with mammoth tusks.

They were so odd that for a long time their evolutionary relationships were unknown. They were generally accepted to be lophotrochozoans, but some studies considered them to be part of their own unique phylum while others tended to place them as being closely related to molluscs.

It wasn’t until 2017 that well-preserved soft tissue fossils revealed a tentacled feeding structure that resembled a lophophore – and hyoliths finally found their place in the lophotrochozoan family tree as close relatives of brachiopods and horseshoe worms, possibly even being a stem lineage within the brachiopod phylum.

However, this isn’t universally accepted and some recent studies continue to dispute it. The feeding organ of a different hyolith fossil has been interpreted as not being a lophophore, classifying the group as an early lophotrochozoan stem lineage, while an analysis of shell microstructure has instead suggested realigning them with molluscs. I’m grouping them with brachiopods here, but future discoveries might still make this obsolete.

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Cambrian Explosion Month #29: Phylum Phoronida & Early Brachiozoans

The last group we’re looking at this month are the brachiozoans, a lineage that includes modern horseshoe worms and brachiopods along with the extinct hyoliths.

Horseshoe worms, or phoronids, are represented by about 15 living species and are usually considered to be their own phylum, but some analyses classify them as a sub-group of brachiopods instead. Like other lophophorates they have a “crown” of filter-feeding tentacles around their mouths, and similarly to some bryozoans they build protective chitinous tubes around their bodies.

There are no definite body fossils of phoronids at all, although there are a few possible trace fossils of their tubes and the enigmatic fossil hederelloids might be related to them.

But some Cambrian fossils might give us a hint about their evolutionary history.

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Cambrian Explosion Month #28: Phylum …Entoprocta?

The only Cambrian fossil species that seems to be closely related to the entoprocts is Cotyledion, but there are several other enigmatic animals that have also been tentatively allied with the group as members of early stem lineages.

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Cambrian Explosion Month #27: Phylum Ectoprocta & Phylum Entoprocta

Ectoprocts, common known as bryozoans or “moss animals”, are aquatic lophotrochozoans that usually live in colonies made up of many tiny cloned zooids. The exoskeletons they build for their colonies have a range of forms, including gelatinous blobs, chitinous branches, and calcified sheets and coral-like fronds.

They’re part of a sub-group of lophotrochozoans called lophophorates, closely related to brachiopods and horseshoe worms, and are characterized by having a ring or U-shaped “crown” of filter-feeding tentacles around their mouths.

Mineralized bryozoans have an extensive fossil record going back to the early Ordovician, about 481 million years ago, but they’re surprisingly absent from the Cambrian – with one possible exception.

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Cambrian Explosion Month #26: Phylum Mollusca – Tentacle Time

Cephalopods‘ highly distinctive body plan and incredible intelligence make them some of the most charismatic marine animals. Today they’re mainly represented by the soft-bodied coleoids (octopuses, squid, and cuttlefish), with the modern giant squid and colossal squid being the largest living invertebrates. In comparison the shelled nautiluses seem like weird oddballs, but they’re actually far more typical examples of the group than their squishier cousins – as part of the conchiferan lineage the ancestors of all modern cephalopods were also shell-bearing molluscs, and for much of their evolutionary history shelled forms like ammonites and orthoceridans were extremely abundant.

The exact evolutionary relationships of cephalopods within the conchiferan family tree aren’t clear, but their closest relatives might be modern monoplacophorans and they probably descended from limpet-like “monoplacophoran-grade” ancestors in the early Cambrian. The current oldest potential cephalopod fossils come from about 522 million years ago, but the first definite cephalopods in the fossil record come from much later in the period.

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