Freshwater squid?

Interesting point about intertidal areas. One thing of note however is that estuarine waters are invariably saline, albeit in fluctuating concentrations. In light of this, it is very likely that cephs in these environments will never actually encounter purely freshwater areas, at least for any length of time. I wonder if osmoregulation, or maybe another problem brought on by salinity levels (bouyancy, possibly?) is to blame? Granted, cephalopods are very similar to gastropods, but given that they fall under the phylum Mollusca, which is more varied than even our own phylum, Chordata, I believe the divide between gastropods and cephalopods is great enough for difference in physiological adaptations.

Of course it could just be entirely down to something as simple as niche allocation. It is entirely possible that any niche the cephalopod could fit in freshwater environments is already taken up by another animal, thus ousting the ceph from filling the niche.
 
The gastropod bit was a stretch, but in my defense, gastropods and cephalopods are (relatively) closely allied within Mollusca. Obviously, since gastropods have invaded those evironments and cephs haven't, they are enough different to for those differences to be meaningful (is that sentence just as clear as mud?). Physiological, however, gastropods and cephs are very similar and if you want to compare cephs to something else physiologically, gastropods are it. So I think it is informative to look for small differences between the two for this question.

I do like you idea about niche competition, but really off the top of my head it would seem the incredibly higher diversity of forms in marine habitats would mean that niches would be all that more closed in marine than freshwater. Freshwater is filled with a few generalist species occupying a wide range on niches that often are occupied by ten times the number of specialist species in the ocean. Also, freshwater habitats is very ephemeral in the evolutionary sense, rapid changes are often leaving large niches open. Then again these quick changes might be part of the problem with ceph invasion. But that answer just beings us back to our original question: other animals do it, why not cephs.

I have on more idea here. Perhaps it is the lack of diversity in the food base in the freshwater that is the problem. Many octopuses have trouble thriving on a monotypic diet... I am not sure I like that explanation either....
 
Taollan;107254 said:
I have on more idea here. Perhaps it is the lack of diversity in the food base in the freshwater that is the problem. Many octopuses have trouble thriving on a monotypic diet... I am not sure I like that explanation either....

Many of us have fed our cephalopods fresh water shrimp, so it is possible for them to prey upon fresh water species.

I am wondering if it has something to do with their high metabolism.

I find it interesting that mammals like the Baikal Seal and the freshwater dolphin species have survived in freshwater environments (although they are now threatened by human pollution...).
 
cuttlegirl;107258 said:
I am wondering if it has something to do with their high metabolism.

There are freshwater fishes that have comparably high metabolism. So I don't think that is the case. What you brought up about freshwater dolphins and Baikals seals is very interesting. Being warm-blooded mammals, they have a much higher metabolism per body weight than octopuses, and thus have a higher food requirement as well. However, since there is relatively little water exchanged through the skin with mammals, osmoregulation is going to be wildly different than with most invertebrates such as octos...
 
cuttlegirl;107258 said:
Many of us have fed our cephalopods fresh water shrimp, so it is possible for them to prey upon fresh water species.

I am wondering if it has something to do with their high metabolism.

I find it interesting that mammals like the Baikal Seal and the freshwater dolphin species have survived in freshwater environments (although they are now threatened by human pollution...).

I like the re-awakening of this thread!

It seems like marine mammals have a rather different approach to osmotic regulation than many other marine animals, in that they tend to keep the land-evolved systems isolated from the water as much as possible, while gill-using animals tend to, by necessity, have a lot of membrane exposure to salt water directly. I'm not sure how marine mammals get hydrated, though.

The fact that so many octopuses live in intertidal zones sure does fly in the face of most of the arguments for why they have never moved into fresh water... another thing that no one has mentioned recently is that cephs are a very ancient group, and I know that the ocean's salinity is believed to have been appreciably less in the Cambrian, so there's some evidence that early shelled cephs existed in lower salinity conditions.

Has anyone else read Peter Ward's Out of Thin Air book? It's overall about how oxygen levels have been a major and often overlooked driving force for evolution (with many hypotheses that Ward advocates looking at, with appropriate caveats that some may well be wrong... it's a food for thought book.) One of the variants of this story is the hypothesis that cephalopod development of jetting and neutral buoyancy, and hence effective predation, was secondary to development of better respiration: that the mantle inhalation and exhalation through the siphon was to get better circulation of low-oxygen water past the gills, and that had the secondary effect of jet-swimming and higher possible metabolism, which offered an opportunity to prey on the oxygen-starved slower critters in the environment, which nudged the evolution in the direction of free swimming, improved jetting, neutral buoyancy, and tentacles to grab lunch. He also suspects the segmentation of trilobites was more about repeating the body segment that has the gill than anything.

I had already been thinking that the presence of so many cephs in deep, low-oxygen water makes many of the "traditional views" of hemocyanin as inferior to haemoglobin is vertebrate arrogance, particularly in light of Gilly's lab's demonstrations of active, fast, huge humbodts spending a lot of time in the anoxic layer, and reports of nautilus having a good ability to survive low-oxygen and even out-of-water conditions. Ward's book, however, really got me thinking a lot more about the inconsistencies of things I'd read, like "the teleost fishes were able to drive the cephalopods to deeper water because their metabolism was much more effective due to haemoglobin and better oxygen storage in tissues." I'm sure this is true up to a point, but the fact that the cephs are the ones who were more successful in the low-oxygen depths suggests that rather than being metabolically challenged, the cephs outcompete vertebrates more readily at low oxygen levels, so the vertebrate respiratory and metabolic systems may be more tuned to current atmospheric oxygen levels rather than universally "better."

I suspect that cephs' mechanisms for handling low oxygen levels are more in the domain of "less well understood" than vertebrates' systems, and there's a tendency to assume "not understood" is "inferior." I'm not sure how osmotic regulation occurs in any mollusc, or how freshwater bivalves and gastropods differ from their marine counterparts in mechanisms for maintaining homeostasis in freshwater environments, and likewise for terrestrial gastropods. I note that table salt is lethal for slugs, snails, and leeches. Is vastly over-salted water lethal to marine molluscs in a similar way? How about freshwater molluscs in marine salinity levels?

Asking why there are no freshwater cephs seems to also just be a reminder of the huge gaps in our understanding of the history of the modern cephs: the almost complete lack of soft body fossils, and consequently our lack of understanding of the prevalence of non-shelled cephs at various points in their evolutionary history-- which raises a question of whether it's easier, harder, or the same for a shelled ceph to migrate to freshwater (where the pumps in the siphuncle and chamber fluid composition might be impacted) or for a coleoid (where it's harder to be neutrally buoyant in freshwater than denser salt water, although octos, for example, seem happy enough to be negatively buoyant.)

I'm not really buying the "all the niches were occupied" argument as an absolute argument, although it may be sort of a "luck of the draw" argument... It seems like cephs were dominant predators in the ocean before vertebrates had established themselves anywhere, certainly not on land. I'd assume that plants and arthropods, and maybe gastropods, had to be well established on land well before vertebrates... if anything, all the hungry ammonites probably provided pressure for arthropods to escape to dry land. Anyone know if there's a fossil record of when the first terrestrial or freshwater gastropods emerged?

A related question someone asked me that may appeal to the readers of this thread is "how come the queen/drone/worker/hive arthropod model that's so successful for insects on land doesn't seem to occur in marine arthropods?" unless there are examples in marine arthropods that I'm unaware of...

And as long as we're asking such questions, perhaps considering why there are no terrestrial or freshwater echinoderms would be interesting as well... I'd think a starfish or sea urchin would be at least as good a candidate as a snail for crawling about on dry land and eating plants and lichens and such off of rocks.
 
Just one of a few good threads on this subject, the following quote from an off topic thread:

Architeuthoceras;3444 said:
Not alot of detail in the following reference, but it does infer that they are rethinking ammonoid habitat and including superhaline, subhaline, and even brackish (surface) water environments. Still not fresh, but ammonoids could have been headed up stream until their demise at the end of the Cretaceous.


(Reference)
Westermann G.E.G., 1996. Ammonoid Life and Habitat. IN Landman, Neil H., Kazushige Tanabe, and Richard Arnold Davis, editors. Ammonoid paleobiology. Plenum Press, New York [ISBN: 0-306-45222-7]

:ammonite:

Also Fresh Water a very similar thread

and a Poll: Are fresh water octos just a hoax?
 
WOO HOO. I love all this sort of discussion...

monty;107263 said:
And as long as we're asking such questions, perhaps considering why there are no terrestrial or freshwater echinoderms would be interesting as well... I'd think a starfish or sea urchin would be at least as good a candidate as a snail for crawling about on dry land and eating plants and lichens and such off of rocks.

As for echinoderms I have a feeling that this is closely tied to their water-vascular system. One of the very basic parts of their lifestyle is this system that keeps a reservoir of water inside them for movement, and this is constantly taking in water from the surroundings at a high volume and leaking water out. On land this would be impossible to maintain (sea star out of water gradually lose the ability to move). This might be feasible in freshwater, but with such a large volume of environmental water inside of you, your osmoregulation problems have gotten to be much much much harder. That is my thoughts on that part..

monty;107263 said:
Has anyone else read Peter Ward's Out of Thin Air book?...I had already been thinking that the presence of so many cephs in deep, low-oxygen water makes many of the "traditional views" of hemocyanin as inferior to haemoglobin is vertebrate arrogance.

No I haven't read that book, but I have really been meaning to. I did talk with him briefly about his ideas on this a year or two ago at the American Malacological Society meetings and I must say that I was very interested.
And I agree...I don't think hemocyanin is as inferior as the traditional view suggests. Octopuses are efficient and extracting O2 from their environment, period. Octopus vulgaris has been shown to be able to take 76% of the oxygen out of each "breath" of water, that is better than humans, and I have found that O. rubescens can regulate their oxygen uptake (keep their oxygen uptake rate the same) down to 7 mmHg (for reference, oxygen saturation at this temp, 11 C, is about 156 mmHg). These numbers are actually superior to most vertebrates. Hemocyanin, however, is only half the equation: Biomechanics plays a large part as well. Octopus respiration is such that water flows over the gills through out the entire ventilatory stroke in one direction. Its not a tidal breath like humans perform, its actually unidirectional, which helps alot. This one direction that water flows is countercurrent to the blood flow in the gill capillary bed, so extraction is additionally efficient. The strategy of having three hearts (two before the gills and one post-gill systemic heart), means they have high blood pressure throughout their arterial system, further helping gas exchange. Also, in terms of hemocyanin, from what I understand it is more efficient at carrying oxygen at low O2 levels. Hemoglobin binds oxygen cooperatively, which means that as you bind more oxygen (up to four per tetramer molecule in humans) it can bind the next more efficiently. This means that if you have well-loaded hemoglobin, it is very effiecient, but the opposite is also true, if there isn't much oxygen, you lose cooperativity and it is no longer that efficient at carrying oxygen, and hemocyanin which show no (or atleast much less) cooperativity keep up relatively the same efficiency as at high O2 levels.
Even in many high-oxygenated near-shore environments cephs seem to hold their own pretty well against vertebrates. I don't know of anywhere cephs are really "rare" in near shore environments. Really, ceph respiratory physiology is very very interesting and entirely under studied. Its much sexier to study behavior and intelligence, but really their respiratory physiology is just as head and shoulders above all other inverts.

monty;107263 said:
A related question someone asked me that may appeal to the readers of this thread is "how come the queen/drone/worker/hive arthropod model that's so successful for insects on land doesn't seem to occur in marine arthropods?" unless there are examples in marine arthropods that I'm unaware of....

As far as I know, even in terrestrial arthropods I think the high social strategy is restricted to hexapoda (the insects), which have extremely few, if any, representatives in marine environments. Terrestrial arachnids, crustaceans, and myriapods are just as asocial as their marine relatives, so maybe the better question isn't marine vs. terrestrial but really hexapods vs. the rest arthropods why this hasn't arisen. And since we are asking question, why are insects, who have been insanely successful in terrestrial and freshwater environments not further invaded marine environments. If they arose on land, I think I could go for the niche exclusion argument on that one... but serious, I really don't know.
 
To address some of the answers also brought up in the other freshwater thread...

And I don't want this to come off as though I am picking on Fujisawas Sake, (s)he simply was the main person proffering ideas, so most of the novel stuff came from them.

Fujisawas Sake;47313 said:
Hemocyanin serves as a good transport, but has a low affinity for, oxygen. Freshwater is more dynamic in the O2 and salinity (ionic) changes than seawater. Fish, utilizing the more oxygen-efficient hemoglobin, were able to colonize freshwater first, and therefore were able to keep this niche well-stocked and defended.

But really this hasn't stopped many other animals that use hemocyanin. And really hemocyanin is good for variable oxygen levels because of reduced cooperativity. Hemoglobin is simply much more efficient at high O2 levels.

Fujisawas Sake;47313 said:
Fish, utilizing the more oxygen-efficient hemoglobin, were able to colonize freshwater first, and therefore were able to keep this niche well-stocked and defended.

But Cephs were around since the upper Cambrian at least, and fish only since the Devonian, it seems cephs had a huge head start and it should have been up to the fish to fight them out of it.

Fujisawas Sake;47351 said:
Maybe a less specialized prey menu might have helped too.

Octopus seem to eat anything that moves, and can get into just about any packaging (clams, finding things in holes and cracks with their arms). Fish tend to be awfully picky eaters, and need soft bodied prey or insects. As for being generally out-competed, it seems an uncomfortable reason for not even seeing a single freshwater ceph anywhere.

Fujisawas Sake;47358 said:
Case in point: octos hunt crustaceans mostly. The majority of the crustacea are still marine. There would have been no need to try new hunting grounds if no need presented itself. The behaviour wouldn't change, so no selective pressure, therefore no evolution toward a freshwater existence.

While there is far fewer species of crustacean in freshwater, there isn't much lack in numbers. Amphipods are numerous (which are often used as prey by juvenile octos) and crayfish occur in high numbers nearly every long-term body of freshwater. And again, octos are one of nature's great dietary generalists. They can eat just about anything fleshy in a pinch (stomache contents have revealed sea cucumbers, gulls, other things that we often think of as unpalatable to these critters.
Secondly, from what I can tell, life will expand until something constrains it. “No need” isn't really a good argument for why X animal doesn't occur in Y location.
Life operates by producing more offspring than can survive. Because there always is a struggle for existence, every possible means of survival will be exploited.

Fujisawas Sake;47358 said:
Another point; osmoregulation. Cephs don't handle freshwater. They are by their nature isosmotic with the seawater and freshwater would stun and most likely kill a ceph quickly.

Kinda circular reasoning. Essentially any marine organism can't handle freshwater (with a few exceptions). If you tossed a marine snail in freshwater it would die, but there are still freshwater snails.
 
monty;107263 said:
I like the re-awakening of this thread!

It seems like marine mammals have a rather different approach to osmotic regulation than many other marine animals, in that they tend to keep the land-evolved systems isolated from the water as much as possible, while gill-using animals tend to, by necessity, have a lot of membrane exposure to salt water directly. I'm not sure how marine mammals get hydrated, though.

Marine mammals get their fluid from the fish/cephalopods that they consume. That being said, when I cared for sick pinnipeds, we would offer them fresh water, and they would readily drink water from a bowl. Also, we would often let them swim in fresh water pools, with no (perceived) ill effect. However, if we were rehabilitating a dolphin or whale, we had to use salt water.
 
monty;107263 said:
I note that table salt is lethal for slugs, snails, and leeches. Is vastly over-salted water lethal to marine molluscs in a similar way? How about freshwater molluscs in marine salinity levels?

For slugs and land snails, I am pretty sure this has to do with being a land animal and osmosis occurring through their skin.

monty;107263 said:
Asking why there are no freshwater cephs seems to also just be a reminder of the huge gaps in our understanding of the history of the modern cephs: the almost complete lack of soft body fossils, and consequently our lack of understanding of the prevalence of non-shelled cephs at various points in their evolutionary history-- which raises a question of whether it's easier, harder, or the same for a shelled ceph to migrate to freshwater (where the pumps in the siphuncle and chamber fluid composition might be impacted) or for a coleoid (where it's harder to be neutrally buoyant in freshwater than denser salt water, although octos, for example, seem happy enough to be negatively buoyant.)

I am wondering whether buoyancy plays some role in this - although I would guess, if a cephalopod were to make the transition to fresh water, it would be an octopus.

monty;107263 said:
And as long as we're asking such questions, perhaps considering why there are no terrestrial or freshwater echinoderms would be interesting as well... I'd think a starfish or sea urchin would be at least as good a candidate as a snail for crawling about on dry land and eating plants and lichens and such off of rocks.

I can't remember the answer to this, but at one point in my academic career I learned the reason there are no freshwater echinoderms, but I think it has something to do with their water vascular system.

Fujisawas Sake;47358 said:
Another point; osmoregulation. Cephs don't handle freshwater. They are by their nature isosmotic with the seawater and freshwater would stun and most likely kill a ceph quickly. Their metanephridia are no where near as hardy as our kidneys, which are pretty fragile. Fish are better at taking the extremes of freshwater by highly developed kidneys. Very few molluscs are freshwater, and those that are show signs of having returned to the water from a land-based life (pulmonate gastropods).

John brings up a good point about cephalopod "kidneys" - their metanephridia may not be capable of handling osmoregulation in fresh water.

For an interesting discussion on the origin of Baikal seals, see this...
Darren Naish: Tetrapod Zoology: The most inconvenient seal


This is a great discussion, if I weren't so sleep deprived, I would be able to respond more intelligently...
 
monty;107263 said:
I note that table salt is lethal for slugs, snails, and leeches. Is vastly over-salted water lethal to marine molluscs in a similar way? How about freshwater molluscs in marine salinity levels?

Yes and yes. Essentially all aquatic animals (with the exception of some estuarine critter) are sensitive to salinity. Maintaining osmotic balance is about as proximal of a need for animals as breathing is. Nearly all aquatic animals exchange essentially all of their fluids through the skin, and changes in salinity royally screw with maintaining that balance. Even the most euryhaline (saline change tolerant) freshwater fish can't survive in marine environments for more than a few minutes, and visa versa. Anadromous fish spend days in estuaries retooling their osmotic physiology before they can make the transition.
Marine mammals are kinda the exception. Having come from land mammals, they retain the water-proof skin, they they are largely resilient to such changes. Every year the in Columbia River sea lions venture far up the river foraging on the migrating salmon, much to the dismay of fisherman and fisheries managers.
So that was a long winded answer to yes, over-salted water will kill just about anything aquatic, including mollusks.
 
But ... Some "guppies" /"mollies" (brackish water fish) CAN and DO survive being simply thrown into salt/fresh water. Granted, a higher success can be achieved by slowly raising/lowering the salinity but survival can occur often enough that any aquarist can accomplish this with long term success without slow acclaimation (acclimitiz(s)ation ;>). Have studies been done on the makeup of the gill "material" of brackish water animals ois there a "pump" difference that would explain the effect?
 
dwhatley;107277 said:
But ... Some "guppies" /"mollies" (brackish water fish) CAN and DO survive being simply thrown into salt/fresh water.

Taollan;107274 said:
Essentially all aquatic animals (with the exception of some estuarine critters) are sensitive to salinity.

Yes, mollies are pretty euryhaline, and can survive in fresh or saltwater because they are a estuarine species. If you have a pretty salinity tolerant animal, adapted to living in estuaries, where salinity is variably on a daily basis, then they are generally somewhat adapted to these changes. Hardy animals like this can generally survive a saline change of about +- 15-17 ppt and survive. So generally anything from 32 ppt (slightly lower than oceanic except Pacific) to 0 ppt is generally tolerated well by estuarine species. That being said, you can kill mollies pretty quickly if you stick them in a stiff brine of 60 ppt or so, and that gets back to my main point of the post:
Taollan;107274 said:
over-salted water will kill just about anything aquatic, including mollusks.
P.S.: I have also found that goldfish are surprisingly tolerant of fairly radical changes in salinity. I have thrown them into 35 ppt saltwater straight from freshwater and had them survive for a couple days. Given, this isn't as good as mollies, but still amazingly good for a non-estuarine fish.
 
Next time we have a student wanting to do research on octopuses, someone remember to point to this thread. Specifically, I am thinking about longevity and hydrating diet ...
 

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