lophotrochozoa – Page 2 – Nix Illustration

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.

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.

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.

Cambrian Explosion Month #22: Phylum Annelida – Down the Wormhole

Among the various stem-polychaete worms known from the Cambrian, the existence of more modern-style annelids like Pygocirrus hint that the common ancestor of modern forms might have evolved much earlier than previously thought. But this was complicated by the fact that all the known stem-polychaetes seem to have been active crawlers or swimmers, while the oldest modern lineage of polychaetes are burrowers, mostly sedentary, and sometimes tube-dwelling.

So if “crown group” forms that lived buried in the seafloor sediment must have diverged at least as far back as the early Cambrian, where were the fossils of them?

Cambrian Explosion Month #22: Phylum Annelida – Plumes and Tails

Annelids in the Cambrian Period were mainly represented by bristle worm polychaetes, with most known species belonging to early stem lineages. And while more modern-style polychaetes would become abundant during the early Ordovician, the earlier Cambrian forms were still surprisingly diverse.

Cambrian Explosion Month #20: Phylum Annelida – Early Worms

Annelids are a large phylum of segmented worms, found worldwide in marine, freshwater, and terrestrial habitats. Their modern diversity includes bristle worms, tubeworms, earthworms, and leeches, and many burrowing species are hugely ecologically important due to their activities aerating and enriching soil and sediments.

As part of the spiralian lineage they’re considered to be members of a grouping called lophotrochozoans, closely related to animals like molluscs, brachiopods, bryozoans, and ribbon worms. But the exact positions of everything within that evolutionary tree is currently a bit uncertain, with different studies coming up with different answers – annelids might be a basal lineage of lophotrochozoans, or they might be closely related to molluscs, or they might instead be the closest relatives of ribbon worms and flatworms.

Like many other soft-bodied animals annelids have a poor fossil record. The Ediacaran Kimberella and Namacalathus may have been early lophotrochozoans, and the abundant “small shelly fossils” of cloudinids known from the Ediacaran-Cambrian boundary (~541 million years ago) may represent the tubes of early annelids. But otherwise some of the first definite annelid body fossils are bristle worm polychaetes from the early Cambrian.

Cambrian Explosion Month #01

It’s March, and that means it’s time for Cambrian Explosion Month!

Orthrozanclus

Orthrozanclus elongata, from the mid-Cambrian of China. Only about 2cm long (~0.8″), this tiny creature was covered in both long spines and extensive armor – with tile-like scales on its back, overlapping dagger-shaped plates around its sides, and a small shell on its head.

It’s the second species of Orthrozanclus to be discovered, extending the genus’ range about 10 million years older than the Canadian O. reburrus.

Although it would have been a member of the Lophotrochozoa (the group that contains modern molluscs, annelid worms, and brachiopods), its exact evolutionary relationships are still uncertain. It might have been a transitional form between Wiwaxia and the halkieriids, or it could be closer related to the brachiopods.