I'm please to announce the lattest The Cephalopod Page contribution. A Broad Brush History of the Cephalopoda is located at: http://is.dal.ca/~ceph/TCP/evolution.html This article is on the evolution of cephalopods and contains images of both fossil and living cephs. Enjoy.

Thanks James, and thank Dr. Monks, what a great article, I thoroughly enjoyed reading that while I was supposed to be working. :D

:ammonite:

Very good article. Clearly written and informative.

Thanks!

Yes, thanks Dr. James... related to this, as has been stated on the TONMO.com ticker for some time now, we'll have our own Ammonite article to post soon... Phil provided a basic overview for the TONMO.com community, which I greatly appreciate! I just have to get around to posting it... :roll: This week Phil, I promise... :oops: :)

Posted it! :)

Ammonites: A General Overview (http://www.tonmo.com/science/public/ammonites.php)

Two Great articles about fossil cephs in one day :D now I am in heaven. I got to read Phils' at home without my boss looking over my shoulder :D GREAT job Phil!

:ammonite:

Outstanding article! Must study more... :read:

Sushi and Sake,

John

Dr. Wood,

Great article! I noticed though that it seems to hint that the cephalopod lineage is somewhat polyphyletic. I see three distinct lines, but is there any idea or theory on what the "proto-cephalopoda" may have been? Also my invert zoo instructor liked to nickname cephs "siphonopoda" because he believed that the siphon evolved from the foot. Any thoughts?

Years ago there was a concept known as the "Hypothetical Ancestral Mollusc" or H.A.M. theory. I posted a subject last year called "H.A.M. and Legs" which asked about the development of the cephalopod arms from the bauplan of the "general" mollusc. I know that H.A.M. theory has gone the way of the of the Ammonites, but is there any thought on what the "proto-cephs" may have actually looked like, and from what branch these creatures may have sprouted?

Thanks for the great article!

Sushi and Sake,

John

Cool article Phil, looks like a lot of work into went into that. I havent got much time but I wanted to let you know, cheers!

Geoff

Years ago there was a concept known as the "Hypothetical Ancestral Mollusc" or H.A.M. theory. I posted a subject last year called "H.A.M. and Legs" which asked about the development of the cephalopod arms from the bauplan of the "general" mollusc. I know that H.A.M. theory has gone the way of the of the Ammonites, but is there any thought on what the "proto-cephs" may have actually looked like, and from what branch these creatures may have sprouted?



Yo John,

Why has the HAM theory been disregarded these days? Is it because it implies the existence of an animal for which we have no evidence for, or has it gone the way of the dodo due to modern cladistic analyses? I suppose the HAM would imply that all the forms of mollusc originated at a single point in time from a single ancestor. I know that there has been much debate about the origin of molluscs recently with some researchers placing the molluscs in early lineages with ancestral annelids. Much of this work comes from recent advances in molecular biology examining RNA sequences in annelids and molluscs. (Just don’t ask me to explain it).

What on earth was going on in the late Precambrian to the Cambrian ‘explosion’ is totally obscure, relying on trace fossils and enigmatic faunas such as the famous Australian Ediacaran fauna. Even the animals that we know of from this mysterious dark age often defy easy description, weird quilted mattresses that may have been animal or vegetable or something in between and tiny impressions of worms and segmented enigmas that seem to have left no clear descendents.

I think it is generally accepted that the earliest molluscs may well have resembled aplacophora such as the ‘living fossil’ Neopilina that was dredged from the ocean floor in the 1950’s. That there were such creatures in evidence in the late Precambrian is highly likely. Scrape marks on rocks are recorded from the late Precambrian period that may well have caused by such primitive molluscs and tiny cap-like shells (a couple of millimetres across in most cases) have been recorded from the Tommotian faunas of Siberia, datable to about 550mya, at the beginning of the Cambrian. Some of these resemble monoplacophora and some bear a similarity to the earliest cephalopods in the late Cambrian such as Palaeoceras and Plectronoceras, though it is clear that they were not cephalopods as they do not have chambered shells containing a siphuncle.

I always imagined that the arms/tentacles on the head of these late Cambrian primitive cephalopods developed from sensitive patches on the head of the primitive proto-cephalopods somewhere in the late Precambrian or early Cambrian, these must have been devices used to feel the immediate local environment, possibly used as a survival aid. Indeed, I imagine they may well have predated the evolution of eyes in molluscs. Perhaps these feelers later developed the secondary function of ensnaring prey when the primitive arms were developed enough to grasp whatever they came across in the environment and push it towards the radula. Thus the path towards a predaceous life amongst cephalopods was set at the earliest days of their ancestry.

I know this is all speculation of mine, and I have never studied zoology, but it sort of makes sense to me!

Cheers,

Phil

(Almost glad he’s off sick for the day; it gave me time to write this!)


BTW, Cheers, GeoffC!

Phil,

That's EXACTLY what they thought the H.A.M. might have looked like! Aplacophoran design is pretty ancestral for molluscs, and it would make sense. My opininon is that they scrapped the theory because there was no solid proof on the H.A.M and not a lot of fossil evidence either way. I still think it bears a second look.

I think it has to do with the idea that scientific theories have to be based on evidence. H.A.M. probably fell by the wayside due to this concept. The problem is, evolution is not always cut and dry, and sometimes a well-educated guess is a good place to start. I think that, given current knowledge of malacological evolution, we can build a backwards model of ceph evolution and be pretty cloes to the truth.

For someone who never studied zoology, you can sure hold your own! 8)

Sushi and Sake,

John

Phil,

I think that there's something brewing as far as deep ocean ecology is concerned... Kat seemed to indicate that there are more species of giant squid out there, and that got my attention. Any thoughts on the ecology of the nautiloids? I wonder how the soft-bodied cephs did during that time?

John

John,

Yes, I think you maybe right.....if I remember correctly, wasn't there a hint on these pages a few weeks ago that there may be another giant out there apart from Archi and Messie? Or perhaps I just misinterpreted a comment (as I often do).

Well, as to your other point, you would need someone much wiser than me to explain the intricacies of ecology of the Palaeozoic oceans. However, if you are referring to 'soft-bodied cephs' as squid, octopi and cuttlefish then there was no real overlap with the nautiloids. The nautiloids really came to dominance way back in the Ordovician period where they certainly assumed the position of top predator. There was an evolutionary explosion of nautiloids at this time with no less than nine orders, most of these had cone shaped or even straight shells and were really quite diverse in their morphologies. Most of these nautiloid groups became extinct by the end of the Permian excepting just two groups, the Orthoceratina which lingered on to the end of the Triassic and, of course, the nautilida from which our modern nautilus is descended. The modern Nautilus really is the sole surviving twig on what was once quite an exotic bush.

The gradual extinction of the nautiloids at the end of the Devonian extending in the case of some orders into the Carboniferous can (probably) partly be explained by the emergence of the ammonoidea in the Devonian, especially the goniatites. Why these should have come to dominate is a very good question, perhaps the ammonoidea were specialists and more adaptable or perhaps they had a faster growth rate and a shorter lifespan so that they could evolve quicker than the nautiloids. If anyone has any good theories, I'd love to hear them here!

Some of the nautiloids were adapted to deep water, indeed, one researcher, Westermann (1985), has established crush depths for these ancient creatures. It is estimated that one nautiloid, the Carboniferous Michelinoceras, could probably withstand depths of 1125m. It seems that direct competition with the shallow-water ammonoids could not be sole reason responsible for explaining the demise of the nautiloids, there must have been other factors as well.

Anyway, going back to your origin point about soft-bodied cephs, the squid and octopi did not appear until the Jurassic era, by which time the only surviving nautiloid order, the Nautilida, had adopted the familiar form we have today and were probably already established in the deep-water niche they occupy today. So it seems likely that there really would not have been that much competition between the two groups. As an aside, I think the cuttlefish are first known from the Cretaceous and as shallow water creatures would certainly not have made much of a direct impact on the Nautilida.

:goofysca: Anyone still awake? :sleeping:

I wonder if their specialization led to their downfall? I mean, the oceans during that period went through large scale changes in relatively short periods of time. Changes would have occoured in salinty, prey distrubution, and in the nature and size of their predators. Even then, I wonder why more of them still aren't around.

As for what I was thinking about giant squid, Kat gave me a resounding "yes" when I asked if she thought there were any more massive squid species out there. *sigh* Better start thinking up more imaginative common names, I guess.... :lol:

Thanks for the reply again! As always, a pleasure to read.

Sushi and Guiness, :beer:

John

Posted it! :)

Ammonites: A General Overview (http://www.tonmo.com/science/public/ammonites.php)

Well done! I'll have to link to that when I have some time to work on TCP.

I think we date ourselves with these theories. I was also taught the HAM theory, that cephalopods evolved from an aplacophora/monoplacophoran ancestor. One of the advantages of this theory was it’s use in teaching as it makes intuitive sense. I’m not sure what happened to it, perhaps it is another “just so” story. Looking at my old invertebrate text book, there was little evidence presented. Modern molecular evidence should tease out the relationships between the various mollusk groups which would help to support or refute this theory. I focus on living cephalopods and am honestly not current in this area.

Dr. Neale Monks wrote the The Cephalopod Page article that kicked off this thread. He is much more knowledgeable about the cephalopod fossil record and their evolution than I am. I’m sure he would be happy to answer questions. I'll send him a copy of this.

Dr. Wood,

See, I don't think H.A.M. is completely wrong, its just that its only a guess, albeit an educated one. I think that the relationship between ancestral forms of molluscs and the cephs is a less direct one, given the overall extreme modification of the body plan.

Think about it like this: take your average rorqual (I went whale-watching today, so I'm still stoked about seeing a gray whale up close) and a human. The skull bones are pretty much the same, but its like they've been re-shaped and pushed and ... well, I'm sure you've taken mammalogy or nat. history of the vertebrates, so you get the idea. We can see the similarities, but overall the body plan of the ceph is so different that I don't see a very direct H.A.M - ceph link. ALTHOUGH... Maybe a pteropod-ceph link may be forseeable. *shrug*

Sorry, that's the cladist in me speaking... :oops:

How accurate do you think that molecular phylogeny will be on such an old phylum? I mean, are they scanning mitochondrial DNA or HOX genes or something else? Taking 500 million years of evolution and mutation, I wonder how exact we could actually be?

Thanks for the reply!

Sushi and Sake,

John

John,

I thought you might be interested to see this.

A 425 million year old creature known as Acaenoplax has been reconstructed in 3D. It comes from a Hertfordshire in the UK and comes from a quite remarkable collection of microfossils that were preserved in Silurian-age nodules. This creature, Acaenoplax is thought to lie somewhere between the aplacophora and polyplacophora. As such it is thought to be a primitive mollusc and therefore related to the earliest ancestors of the cephalopods. (Though, of course, cephalopods were already established as a group in their own right by that date).

Strangely it lacks the classic molluscan foot but looks like a spiny caterpillar.

Acaenoplax; an early mollusc (http://www.earth.ox.ac.uk/herefordshire/acaen/acaenoplax.htm)

Phil,

Dude, that was brilliant! Thanks for the heads up.

I was thinking about this subject the other day, and I think the discovery of Acaenoplax lends some credence to the annelid-mollusc link. It looks like a polychaete, sans really noticeable evidence of metamerism in its bauplan (body plan). I don’t think this is accidental though. A little background on why:

In the book Invertebrates, by Steve and Gary Brusca (1990), four individual theories of molluscan evolution are presented. They are:

The Turbellarian Theory - This theory states that molluscs may have evolved from acoelomate turbellarians, and it based on the homologous (or analogous) mucociliary gliding surface. They don’t like this theory, and I think it’s a long shot at best. I'm sorry, but I'm just not that much of a cladist.

The Modified-Turbellarian Theory – This theory states that molluscs evolved alongside annelids from a turbellarian ancestral group. Hmm… not sure there either. This seems to be dependent on the idea that the aceolmate design "branched" into the annelid coelom and molluscan hemocoel. I think this is also a weak theory, and I go into a bit more detail below.

The Coelomate Theory – This theory states that molluscs evolved from a coelomate ancestor alongside the annelids. Given the idea that molluscs and annelids both show embryonic spiral cleavage, 4d mesentoblast development, and trochophore larvae, I think this is a good theory.

And last, but not least, The Annelid Theory – This theory states that molluscs are direct descendants of annelids. The Bruscas go on to say that the major problem with this theory is the apparent “de-evolution” of segmentation and coelomic development that would have had to occur in the annelida. Given the aplacophorans and caudofoveatans, and their apparent modifications of the molluscan bauplan, I’m not entirely sold that this isn’t the case. I’ll explain below.

For one thing, “de-evolution” is not an exact term; indeed, body plans are highly modified to suit new patterns in genetic expressions, environmental adaptations, etc. Good examples are arthropod parasites, most notably parasitic crustaceans. Remember the rhizocephalan? At first glance, it appears to be a fungal endo-parasite, assuming hormonal and developmental control of its unlucky crab host. But its larval form betrays its true ancestry. It has a cyprid larva; it is a cirripedian… a barnacle. The modifications on that body plan are insane. I think it was on a neotenic path, and then somehow took a parasitic turn to the dark side. A “simple” parasitic body plan is the result of millions of years of evolutionary specialization, and therefore is not “simple” at all.

The caudofoveatans and aplacophorans are given the “ancestral”, or “primitive” moniker due to their shell-less bodies. Cladists tend to put them low on the molluscan cladogram. Remember H.A.M.? Well, these two groups are enigmatic because, if these cladists are right, the molluscan shell would have vanished from a shelled H.A.M. ancestor, then reappeared. The book says that this is unlikely. HOWEVER, a recent finding in the insecta (Published in Scientific American) seems to indicate that body parts “lost” in the evolutionary process can return (in the given case, it was wings). Darn, I need to find that article… If this is true, then the gene or genes for shell formation would still be found in these two classes, and the lack of shell would be due to extensive modification of their molluscan bauplan.

Imagine evolution reversing its own engineering! Stranger things have happened…

Annelids are segmented, and eucoelomate, but each body part has subtle variations that make it unique. Given their protostome heritage, it is not entirely inconceivable that one or more groups would gradually modify the existing body plan in such a way that the coelom would also be affected. Over time, an ancestral form of annelid (shelled polychaete maybe) could have begun a move toward a less segmented, more molluscan, open-hemocoel form. This is not truly a “de-evolution”, rather it’s a shift in development due to selective pressures and expression of changes in linked genes.

And given the developmental difference, yet structural similarity between coeloms in protostomes and deuterostomes, I think that a lot of genetic analysis will need to be done before we can even consider the coelom as an evolutionary link between groups. The coelom may just be a great idea run through the convergent evolution copy machine.

Hmm... I do play the polyphyly card a lot, don’t I? :heee: Hell, maybe its just my way of copping out, but I can't shake the feeling that similar characteristics do not homologues make.

If the annelid-mollusc links are true, Acaenoplax could be the “missing link” between molluscs and annelids. This is definitely worth a second look.

AUGH!! WHERE IS THE PROOF?!?! @$#%&$*

*sigh* The truth is out there.... oops, wait... that's the X-Files...

Sorry about being so long winded… I think I would have been a lot more coherent if it weren’t for the insomnia.

Sushi and Sake,

John

HOWEVER, a recent finding in the insecta (Published in Scientific American) seems to indicate that body parts “lost” in the evolutionary process can return (in the given case, it was wings). Darn, I need to find that article…

FOUND IT!!!!

http://magma.nationalgeographic.com/ngm/0309/resources_geo3.html

Sushi and Sake,

John

Sometimes, one just HAS to ressurect old threads, and this one is an oldie but goodie...

Any new information on phylogeny of molluscs out there?

John

Thanks to Phil & Dr. Wood for taking the time to create 2 great articles. Certainly filled in some yawning gaps & clarified a few points I've struggled with :read:
I feel like my "party balloon" knowledge base has been connected directly to the Zeppelin like same of these two fine cephalopodan pedagogues & I'm fillin' up ! :notworth:
After a Bank holiday weekend having brushed up on the Cambrian, gained some invaluable prepping experience & read these 2 articles I'll confidently take on a room full of Eastenders viewers at PaleoTriv !! :grad:

"Although not as glamorous as Tyrannosaurus rex or as dramatic as Velociraptor, these extinct creatures are nonetheless quite interesting animals."

Fascinating as they are in their own right, in my opinion the creatures that have & still inhabit the oceans are light years ahead in the glamour & drama dept. of any of us landlubbers i.e
Tyrannosaurus rex v. Liopleurodon ferox ? no contest :whalevsa:

Hi John, Spartacus (Hey, I'M SPARTACUS!!!),

Any new information on phylogeny of molluscs out there?

To be honest John, I really have not looked that sort of stuff up for many months so I'm afraid I couldn't really say. If I do find anything of interest, I'll let you know. Sorry, old chap!

After a Bank holiday weekend having brushed up on the Cambrian, gained some invaluable prepping experience & read these 2 articles I'll confidently take on a room full of Eastenders viewers at PaleoTriv !!

Thanks, Spartacus! Actually I was thinking about going back and changing much of that ammonite article one of these days. It was the first one of these things that I wrote and to be honest, does not really talk about the ammonites themselves that much (and the pictures are crap)...I'll get around to it one of these days. I'm trying to pull together something on prehistoric giant squids at the moment, but information is very hard to find. It will be a short one this time.

As for the Cambrian stuff, if you want to read more I would recommend:

Steven J Gould "Wonderful Life" (1989) on the Burgess Shale. A classic, even read by Jimmy Carter!

Simon Conway-Morris "Crucible of Creation"(1999). Dry reading and really a reply to Gould but written by one of the top researchers in the field so worth getting.

Richard Fortey: "Trilobite!" (2001). A brilliant little book which makes trilobites appear quite exciting and is very funny in places and full of amusing anecdotes. Fortey is a senior palaeontologist at the British Museum of Natural History in London yet somehow manages to bring up Bond villains, Australian pubs in the Outback and The Beatles. After you read this, you'll really want to find one yourself.

Also recommended about the evolution of marine life is Richard Ellis' book "Aquagenesis" (2001) which covers practically every major group of marine animals, vertebrate and invertebrate, and is very clearly written with nice illustrations. There's a good section on cephalopods too.

Phil

I'm Spartacus Artisticus, you're welcome & way too modest. Thanks for the recommended reads too, Trilobite sounds right up my street & I could read it to my pet flexi-calymene. There must be something in the water, formaldehyde possibly, at the NHM as I believe they may all be a tad eccentric. I attempted to get my name in neon lights via the entomolgy dept. with the 1st capture of otiorynchus armadillo in Norfolk. All I'd actually subdued was an otiorynchus sulcatus, in my defence they're like peas in a pod & the defining difference being something like the urine dewpoint.
BTW I've a theory (already, for one so novice you all cry) on the demise of your superbly illustrated plectronoceras which I'm surprised all you bright buttons that I now mix with haven't noticed as it IS a major design fault.
Just take a look at how close this poor creature's gills are to its anus !! :shock:
Put yourself in a plectronoceras' shoes for a minute, these creatures weren't wiped by outside influences, they just gave up ! :angel:

Whoa,

Is this the little critter to which you refer?

http://www.rhs.org.uk/publications/pubs/garden1202/newspests.asp

I had no idea that we were under attack by alien weevils in Southern Britain! Mind you, we get all sorts of peculiar invertebrates here in Dover, being a major port. I'm not referring to the contents of the local night club, but peculiar colonies of tiny Mediterranean scorpions infesting the old stone walls in the dock area and parts of the town. They apparantly glow under UV light, though I have not tried it myself (always wondered who first discovered that - strange). Must try and find out exactly what they are one day.

All sorts of horrendous things scuttle off the freight ships docking, especially those carrying fruit and foodstuffs. Most die quickly but not all......Another pub in the town centre that shall remain nameless, has a colony of Black Widows out the back. Had a good look at one close up once, really should have brought a camera.

Good point about the Plectronoceras, its a wonder how any animal that breathed through a cloud of its own detritus could have evolved into anything impressive... :)

that's the fella, grab him !! I'm still convinced I was right & Max at the NHM
is unwilling to share the limelight. He didn't even return my film pot :roll:
Seems they hide in shipments of shrubberies, courtesy of Roger the Shrubber from Italia.

Dover sounds like a bug fan's Utopia. I remember seeing Sir Dave Attenborough waving a UV lamp about down there showing off them wee critters & a few years earlier (lots) he was showing off the (native ?) scorpions of Ongar station.

always wondered who first discovered that - strange
what's stranger is which deviant found cow's milk was yummy !


Dover sounds like a bug fan's Utopia. I remember seeing Sir Dave Attenborough waving a UV lamp about down there showing off them wee critters & a few years earlier (lots) he was showing off the (native ?) scorpions of Ongar station.

always wondered who first discovered that - strange
what's stranger is which deviant found cow's milk was yummy ! :yuck:

its a wonder how any animal that breathed through a cloud of its own detritus could have evolved into anything impressive

There's a lot of people out there who talk detritus !

Time to make like George Romero and raise the dead thread! On Friday, a guest lecturer at Humboldt State University will be speaking on the Green Chiton. The lecture is titled "Green Chiton Eggs and H.A.M." and I hope to get the lowdown on some great mollusc development information and what happened to H.A.M. (Hypothetical Ancestral Mollusc) theory.

Hope to post more soon! Trying to earn my keep as a SC,

John