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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Cambrian Explosion Month #25: Phylum Mollusca – Shelling Out

The exact evolutionary relationships of the main groups of modern molluscs are rather debated, with several different proposed family trees. But one of the main possibilities is that there are two major lineages: the aculiferans and the conchiferans.

Modern conchiferans include slugs and snails, cephalopods, bivalves, tusk shells, and monoplacophorans – all groups that ancestrally have either a single-part shell or a two-part bivalved shell, with some lineages later becoming secondarily shell-less.

The ancestral conchiferans are thought to have been monoplacophoran-like molluscs, limpet-like with a cap-shaped shell, and likely diverged from a common ancestor with the aculiferans around the end of the Ediacaran. (But modern monoplacophorans probably aren’t “living fossil” descendants of early Cambrian conchiferans, and may instead be close relatives of cephalopods that have convergently become similar in appearance to their ancestors.)

Some of the earliest conchiferans were the helcionelloids, a lineage of superficially snail-like molluscs with coiled cone-shaped mineralized shells. They appeared in the fossil record at the start of the Cambrian (~540-530 million years ago) and lasted until the early Ordovician (~480 million years ago), and have been found all around the world as components of the “small shelly fauna“.

And while they’re usually tiny, only a couple of millimeters in size, they may actually represent juveniles or larvae – there’s evidence that at least some species grew up into much larger 2cm (0.8″) limpet-like adult forms.

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Cambrian Explosion Month #24: Phylum Mollusca – Coats of Mail

Much like Odontogriphus and Wiwaxia, the evolutionary relationships of a group called the halkieriids have been debated for a long time. These animals looked like “slugs in chain mail“, covered in thousands of tiny overlapping mineralized armor plates along with a larger shell plate at each end.

In the past they’ve been assigned to different parts of the lophotrochozoan family tree, sometimes being placed closer to annelids or brachiopods, but at this point they’re generally accepted to be molluscs. The spiny species Orthrozanclus may link halkieriids with wiwaxiids in a larger “halwaxiid” lineage of early molluscs – or they might instead be early members of a group called aculiferans

Aculiferans are represented in modern times by chitons and aplacophorans, and they’re distinguished from all other molluscs by having either eight shell valves (chitons) or no shell at all and a worm-like body covered with tiny calcareous spines (aplacophorans).

(Also chitons are especially weird, with magnetite teeth and thousands of eyes in their armor plates.)

A related fossil species called Calvapilosa kroegeri from the early Ordovician of Morocco (~480 million years ago) seems to link halkieriids with aculiferans, placing the chain-mail-slugs as a stem lineage close to the common ancestor of modern forms.

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Cambrian Explosion Month #23: Phylum Mollusca – The Stem Weirdos

Molluscs are one of the largest animal phylums, second only to the arthropods, and are also hugely diverse, found in marine, freshwater, and terrestrial environments all over the world. Not only are familiar modern animals like bivalves, slugs and snails, and squid and octopuses included in this huge lineage, but also nautiloids, chitons, tusk shells, monoplacophorans, worm-like aplacophorans, and the extinct ammonites and orthocerids.

Like the annelids they’re lophotrochozoan spiralians, and their exact evolutionary relationships within that group are a bit uncertain. But their fossil history seems to go back at least 558 million years with the “mollusc-like” Ediacaran Kimberella, and the earliest members of most major mollusc lineages had probably already diverged from each other before the start of the Cambrian.

The common ancestor of all molluscs probably had features like an unsegmented body, a muscular foot on their underside, a mantle and mantle cavity, a radula, and possibly a tough but non-mineralized leathery “shell” – and Odontogriphus omalus may represent an early stem lineage retaining that basic body plan into the mid-Cambrian.

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Cambrian Explosion Month #19: Phylum Chaetognatha

Chaetognaths, commonly known as arrow worms, are a major component of marine planktonic ecosystems all around the world. They’re a fairly small phylum in terms of diversity, with only about 120 known modern species, but in sheer numbers of individuals they’re incredibly abundant – making up as much as 15% of total zooplankton biomass worldwide. They play an important role as predators, feeding on things like copepods, fish larvae, and each other, and can be so voracious that they’re sometimes nicknamed “tigers of the zooplankton”.

And they’ve been doing it for a very long time.

The appearance of protoconodont “teeth” at the start of the Cambrian (~541 million years ago) suggests that arrow-worm-like gnathiferans were some of the first active swimming planktonic predators – taking advantage of ecosystems that were becoming increasingly complex around that time, and laying the early foundations for more modern-style marine food chains.

Unfortunately we don’t know much about their evolutionary origins, with their small fragile soft bodies leaving only a very patchy fossil record. Their relationship to other animals was also rather enigmatic for a long time, and they were only very recently identified as being part of the gnathiferans.

But their ancestors may have been something like Dakorhachis thambus.

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