Air breathing fish again

[Paul Ciszek]

The recent article on living fossil fish mentioned at least one obligate
air breather, and that renewed my curiousity on this subject.

1) Which fish other than the Dipnoi can be said to have lungs?

2) Which fish other than the Dipnoi are obligate air breathers?

3) Is it undisputed that swim bladders are degenerate lungs and
not the other way around?

4) Do coelacanths show any evidence of lungs?

5) Is there another biology newsgroup, not dedicated to arguing about
creationism, where people like to discuss stuff like this? sci.
bio.evolution seems kind of dead.

--
Please reply to: | "Evolution is a theory that accounts
pciszek at panix dot com | for variety, not superiority."
Autoreply has been disabled | -- Joan Pontius

[Ernest Major]

In message <jojcni$h97$1...@reader1.panix.com>, Paul Ciszek
<nos...@nospam.com> writes

>The recent article on living fossil fish mentioned at least one obligate
>air breather, and that renewed my curiousity on this subject.
>
>1) Which fish other than the Dipnoi can be said to have lungs?

At least three of the four non-teleost orders of ray-finned fishes have
lungs - bichirs and reedfishes, gars, and bowfins.

>
>2) Which fish other than the Dipnoi are obligate air breathers?

Not all Dipnoi are obligate air breathers.

>
>3) Is it undisputed that swim bladders are degenerate lungs and
> not the other way around?
>
>4) Do coelacanths show any evidence of lungs?
>
>5) Is there another biology newsgroup, not dedicated to arguing about
> creationism, where people like to discuss stuff like this? sci.
> bio.evolution seems kind of dead.
>

--

Alias Ernest Major

[Richard Norman]

On Fri, 11 May 2012 15:54:26 +0000 (UTC), nos...@nospam.com (Paul
Ciszek) wrote:

>The recent article on living fossil fish mentioned at least one obligate
>air breather, and that renewed my curiousity on this subject.
>
>1) Which fish other than the Dipnoi can be said to have lungs?
>
>2) Which fish other than the Dipnoi are obligate air breathers?
>
>3) Is it undisputed that swim bladders are degenerate lungs and
> not the other way around?
>
>4) Do coelacanths show any evidence of lungs?
>
>5) Is there another biology newsgroup, not dedicated to arguing about
> creationism, where people like to discuss stuff like this? sci.
> bio.evolution seems kind of dead.

1) A non-technical discussion about this is at
Air Breathing in Tropical Freshwater Fishes
http://www.fbas.co.uk/ABOS.html
It says that air breathing is found in 49 families. Not all use
lungs/swim bladders though. There are other organs that function in
respiration.

2) The gourami, Trichogaster trichopterus, is an obligate air
breather. See
http://jeb.biologists.org/content/82/1/215
Aerial CO2 Excretion in the Obligate Air Breathing Fish Trichogaster
Trichopterus: A Role for Carbonic Anhydrase

The pirarucu, Arapaima gigas, is another example

http://www.nrcresearchpress.com/doi/abs/10.1139/z78-136?journalCode=cjz
Carbon dioxide excretion in the pirarucu (Arapaima gigas), an
obligate air-breathing fish

There is a long list in "The Physiology of Tropical Fishes", in the
chapter on "The Cardiorespiratory System", p. 257, but google books
doesn't allow cut and paste.

3) Is anything ever undisputed? There are sufficient differences
between the structure and development of fish lungs and fish swim
bladders to put doubt on their relationship. See, for example,

Which came first, the lung or the breath?
http://www.sciencedirect.com/science/article/pii/S109564330100304X

The abstract says: "Lungs are paired ventral derivatives of the
pharynx posterior to the gills. Their respiratory blood supply is the
sixth branchial artery and the venous outflow enters the heart
separately from systemic and portal blood at the sinus venosus
(Polypteriformes) or the atrium (lungfish), or is delivered to a
separate left atrium (tetrapods). The swimbladder, on the other hand,
is unpaired, and arises dorsally from the posterior pharynx. It is
employed in breathing in Ginglymodi (gars), Halecomorphi (bowfin) and
in basal teleosts. In most cases, its respiratory blood supply is
homologous to that of the lung, but the vein drains to the cardinal
veins. Separate intercardiac channels for oxygenated and deoxygenated
blood are lacking. The question of the homology of lungs and
swimbladders and of breathing mechanisms remains open."


4) Coelocanths do have lungs. The fossil ones have ossified lungs
and the modern ones have fat-filled "pseudo" lungs.

5) The topic "Top 10 'living fossil' fishes" is discussed in
sci.bio.paleontology with a number of t.o. regulars participating
there, including phyikos, John Harshman, Arrkalen, Paul Ciszek, and
UC. It is not at all clear that is a better place for the discussion.

[Arkalen]

On 11/05/12 16:54, Paul Ciszek wrote:
> The recent article on living fossil fish mentioned at least one obligate
> air breather, and that renewed my curiousity on this subject.
>
> 1) Which fish other than the Dipnoi can be said to have lungs?
>
> 2) Which fish other than the Dipnoi are obligate air breathers?
>
> 3) Is it undisputed that swim bladders are degenerate lungs and
> not the other way around?

The word you're looking for is "exapted", not "degenerate" (it makes me
wonder what the "other way around" would be, lungs are degenerate swim
bladders ?).

I am absolutely no expert but I couldn't find any source disputing it.
Then again given primitive lungs and primitive swim bladders are
basically an air sac connected to the gut, it could be that the
ancestral organ was dual-purpose.

>
> 4) Do coelacanths show any evidence of lungs?

Modern Latimeria coelacanths have a fat-filled organ that's probably
homologous to lungs. Older coelacanth fossils seem to have an equivalent
organ except bony and filled with air instead of fat.

[Arkalen]

... Or, you know, I could just have waited for Richard Norman's response
to appear... :p

[drose...@yahoo.com]

On Friday, May 11, 2012 12:57:38 PM UTC-4, Richard Norman wrote:
> On Fri, 11 May 2012 15:54:26 +0000 (UTC), nos...@nospam.com (Paul
> Ciszek) wrote:
>
> >The recent article on living fossil fish mentioned at least one obligate
> >air breather, and that renewed my curiousity on this subject.
> >
> >1) Which fish other than the Dipnoi can be said to have lungs?
> >
> >2) Which fish other than the Dipnoi are obligate air breathers?
> >
> >3) Is it undisputed that swim bladders are degenerate lungs and
> > not the other way around?
> >
> >4) Do coelacanths show any evidence of lungs?
> >
> >5) Is there another biology newsgroup, not dedicated to arguing about
> > creationism, where people like to discuss stuff like this? sci.
> > bio.evolution seems kind of dead.
>
> 1) A non-technical discussion about this is at
> Air Breathing in Tropical Freshwater Fishes
> http://www.fbas.co.uk/ABOS.html
> It says that air breathing is found in 49 families. Not all use
> lungs/swim bladders though. There are other organs that function in
> respiration.
>
> 2) The gourami, Trichogaster trichopterus, is an obligate air
> breather. See
> http://jeb.biologists.org/content/82/1/215
> Aerial CO2 Excretion in the Obligate Air Breathing Fish Trichogaster
> Trichopterus: A Role for Carbonic Anhydrase

The gourami breathes air. However, it doesn't use its swim bladder
to do so. The gourami uses a branchial organ, which is modified gill.
The branchial organ is not homologous to a lung. The gourami does have a
closed swim bladder that is committed to flotation rather than respiration.
Many air breathing fish do not use their swim bladders to breath. If
you mention the gourami as an air breathing fish, then you should mention
some of these others.
The electric eel is an obligate air breather. The electric eel has a
highly vascularized "cheek". It swallows air, which it keeps in its
pharynx. The vascularized cheek absorbs the oxygen. The electric eel
does not use a swim bladder for respiration.
Dissolved carbon dioxide is dispersed in the water through the
electric eels skin. Many obligate air breathers among fish and amphibians
get their oxygen by breathing in air at the surface, but expel dissolved
oxygen through their skin.

> 3) Is anything ever undisputed? There are sufficient differences
> between the structure and development of fish lungs and fish swim
> bladders to put doubt on their relationship. See, for example,
>
> Which came first, the lung or the breath?
> http://www.sciencedirect.com/science/article/pii/S109564330100304X
>
> The abstract says: "Lungs are paired ventral derivatives of the
> pharynx posterior to the gills. Their respiratory blood supply is the
> sixth branchial artery and the venous outflow enters the heart
> separately from systemic and portal blood at the sinus venosus
> (Polypteriformes) or the atrium (lungfish), or is delivered to a
> separate left atrium (tetrapods). The swimbladder, on the other hand,
> is unpaired, and arises dorsally from the posterior pharynx. It is
> employed in breathing in Ginglymodi (gars), Halecomorphi (bowfin) and
> in basal teleosts. In most cases, its respiratory blood supply is
> homologous to that of the lung, but the vein drains to the cardinal
> veins. Separate intercardiac channels for oxygenated and deoxygenated
> blood are lacking. The question of the homology of lungs and
> swimbladders and of breathing mechanisms remains open."

Another hypothesis that I learned on the Internet (!) was that
the air breathing function reevolved in some swim bladders. The
"lung" had become closed in the ancestors gar, carp and catfish.
However, it opened again at some later time to become a lung again.
This hypothesis does not contradict the idea that the swim bladder first
originated as a "lung". Some may claim that Dollo's Law was violated, but
Dollo's Law itself can be questioned. I don't know how many legitimate scientists believe that the air breathing lung had reevolved.
BTW: The goldfish has a "lung". Yes, this domesticated carp has an
open air bladder that it uses when the water in its tank goes stale. On
warm days, you can watch goldfish in a pond come to the surface to gulp
air. Warm water does not dissolve gases like oxygen. So in the summer, when
the pond water is warm, the goldfish becomes an obligate air breather !-)

> 4) Coelocanths do have lungs. The fossil ones have ossified lungs
> and the modern ones have fat-filled "pseudo" lungs.

Interesting.

[Richard Norman]

You are quite correct in emphasizing the use of breathing organs other
than lungs/swim bladders. I just mentioned it in passing and it is
too easily passed by. I interpreted the question on obligate air
breathing technically and narrowly without regard to what organ is
involved. The real question is the relation of tetrapod lungs to fish
lungs and swim bladders.

[jillery]

On Fri, 11 May 2012 15:54:26 +0000 (UTC), nos...@nospam.com (Paul
Ciszek) wrote:

>The recent article on living fossil fish mentioned at least one obligate
>air breather, and that renewed my curiousity on this subject.
>
>1) Which fish other than the Dipnoi can be said to have lungs?
>
>2) Which fish other than the Dipnoi are obligate air breathers?
>
>3) Is it undisputed that swim bladders are degenerate lungs and
> not the other way around?

Is degenerate a synonym for derived?

[jillery]

ISTM comparing embryological development should shed some light on
this question.

[Richard Norman]

On Fri, 11 May 2012 15:35:31 -0400, jillery <69jp...@gmail.com>

wrote:

>On Fri, 11 May 2012 12:57:38 -0400, Richard Norman
><r_s_n...@comcast.net> wrote:

<snip except relation between fish lungs and swim bladders>

>> Is anything ever undisputed? There are sufficient differences
>>between the structure and development of fish lungs and fish swim
>>bladders to put doubt on their relationship. See, for example,
>>
>> Which came first, the lung or the breath?
>> http://www.sciencedirect.com/science/article/pii/S109564330100304X
>>
>>The abstract says: "Lungs are paired ventral derivatives of the
>>pharynx posterior to the gills. Their respiratory blood supply is the
>>sixth branchial artery and the venous outflow enters the heart
>>separately from systemic and portal blood at the sinus venosus
>>(Polypteriformes) or the atrium (lungfish), or is delivered to a
>>separate left atrium (tetrapods). The swimbladder, on the other hand,
>>is unpaired, and arises dorsally from the posterior pharynx. It is
>>employed in breathing in Ginglymodi (gars), Halecomorphi (bowfin) and
>>in basal teleosts. In most cases, its respiratory blood supply is
>>homologous to that of the lung, but the vein drains to the cardinal
>>veins. Separate intercardiac channels for oxygenated and deoxygenated
>>blood are lacking. The question of the homology of lungs and
>>swimbladders and of breathing mechanisms remains open."
>
>
>ISTM comparing embryological development should shed some light on
>this question.
>

See "Deconvoluting lung evolution: from phenotypes to gene regulatory
networks"
http://icb.oxfordjournals.org/content/47/4/601.full

It says, among other things

The hypothesis—based on extant phylogenetic bracketing—that
Osteichthyes originally possessed paired ventro-lateral pharyngeal
pouches that could have given rise to lungs is supported by
embryological data. There is no embryological evidence, however, to
support the migration hypothesis, by which the paired ventral lungs
are presumed to have formed the unpaired, dorsal swimbladder (Moser
1904). The most plausible hypothesis is that both have their origins
in a “respiratory pharynx” (Neumayer 1930; Wassnetzov 1932). Thus, it
is unlikely that lungs and the swimbladder are derived one from the
other, since none of the three homology criteria (same relative
location, same embryological origin, ontogenetic, or phylogenetic
continuum) used in comparative anatomy and evolutionary biology are
satisfied. Even the discovery of activity of the same genes in lung
and swimbladder development would not prove the homology of the
organs: hand and foot are not homologous although there genetic
patterns of development are similar.

[prawnster]

On May 11, 11:23 am, drosen0...@yahoo.com wrote:
> [...]

>     Another hypothesis that I learned on the Internet (!) was that
> the air breathing function reevolved in some swim bladders. The
> "lung" had become closed in the ancestors gar, carp and catfish.
> However, it opened again at some later time to become a lung again.
>     This hypothesis does not contradict the idea that the swim bladder first
> originated as a "lung". Some may claim that Dollo's Law was violated, but
> Dollo's Law itself can be questioned. I don't know how many legitimate scientists believe that the air breathing lung had reevolved.
>     BTW: The goldfish has a "lung". Yes, this domesticated carp has an
> open air bladder that it uses when the water in its tank goes stale. On
> warm days, you can watch goldfish in a pond come to the surface to gulp
> air. Warm water does not dissolve gases like oxygen. So in the summer, when
> the pond water is warm, the goldfish becomes an obligate air breather !-)
>

Is it useful in any way to tell a story about a "lung" closing in the
gar, carp, and catfish and then reopening again at some later time to
become a lung again? Also, why would it ever be important whether
anyone believed that an air breathing lung reevolved? How is any of
this useful? How is any of this evidence that anything ever evolved?
Is evolution just an insular, incestuous, fetishist hobby not much
different from any other pointless hobby, such as memorizing every
line of every Star Wars movie?

If I were sitting in a room and overheard this discussion, I would
slip into a coma out of sheer boredom.

I will designate this discussion officially off-topic.

[RAM]

It's all about you, prawnster the silly troll.

[jillery]

On Fri, 11 May 2012 16:45:05 -0400, Richard Norman

From your cite:

"The swim bladder and lungs of aquatic and terrestrial organisms arose
independently from a common primordial “respiratory pharynx” but not
from each other."

I'm glad to know my lungs aren't degenerate swim bladders, even if I
am a total degenerate.

[jillery]

Hopefully someone will take pity on you and tape a sign saying "DNR"

[Richard Norman]

On Sat, 12 May 2012 01:25:29 -0400, jillery <69jp...@gmail.com>

And the fish (I mean REAL fish, those bony things with ray fins that
swim in water) are equally glad to know their swim bladders aren't
degenerate lungs.

[jillery]

On Sat, 12 May 2012 10:15:21 -0400, Richard Norman

As another aside, several posters to this topic identified several
species of organisms with gills which also have organs, not
necessarily lungs, which let them breathe air. But to the best of my
knowledge, no aquatic tetrapod has any organ to breathe water. Or do
tadpoles qualify?

[Arkalen]

I think they should, but if they don't surely the Axolotl would.

I guess it is telling though that it's so rare. I read somewhere that
oxygen is much easier to get from the air than from water, so aquatic
species that had lungs were better off sticking with that. (of course
this would be more true of species with very advanced lungs and high
metabolic rates like mammals and birds, than of amphibians)

By the way, on the lungs oxygenate the heart in fishes thing, this more
recent article :
http://onlinelibrary.wiley.com/doi/10.1111/j.1095-8649.2007.01496.x/abstract
(hope this link works better than the previous one)

seems to argue against that theory. So it seems the question is still in
the air even after fifteen years.

[Richard Norman]

On Sat, 12 May 2012 13:29:40 -0400, jillery <69jp...@gmail.com>

Modern amphibians of all three orders are typically aquatic in their
larval stage with external gills. Tadpoles are very similar to fish
in many aspects of physiology although fish gills are internal, not
external.

A number of salamanders have gills as adults. The axolotl along with
some other members of the genus Ambystoma, have external gills
although they are more technically neotonic. The mudpuppy, Necturus,
as another that doesn't lose its gills.

Amphibians also use their skin as a major respiratory organ. Adult
bullfrogs can get 80% of their oxygen through their skin. This
property is used both in air and while submerged.

Some turtles can take in oxygen through their skin in a form of
aquatic respiration to extend the period they can remain submerged.

[Richard Norman]

On Sat, 12 May 2012 14:17:04 -0400, Richard Norman

Add sea snakes to the list of aquatic respirers, again using their
skin.

[Ernest Major]

In message <1Wh*eL...@news.chiark.greenend.org.uk>, Arkalen
<ark...@inbox.com> writes

Adult frogs have an organ to breathe water. It's called the skin.

http://en.wikipedia.org/wiki/Frog#Respiration_and_circulation

>
>I guess it is telling though that it's so rare. I read somewhere that
>oxygen is much easier to get from the air than from water, so aquatic
>species that had lungs were better off sticking with that. (of course
>this would be more true of species with very advanced lungs and high
>metabolic rates like mammals and birds, than of amphibians)
>
>By the way, on the lungs oxygenate the heart in fishes thing, this more
>recent article :
>http://onlinelibrary.wiley.com/doi/10.1111/j.1095-8649.2007.01496.x/abst
>ract (hope this link works better than the previous one)
>
>seems to argue against that theory. So it seems the question is still
>in the air even after fifteen years.
>

--
Alias Ernest Major

[drose...@yahoo.com]

> But to the best of my
> knowledge, no aquatic tetrapod has any organ to breathe water.

Adult amphibians generally "breath" through their skin. Their
skin does not have any scales. The skin is moist and covered with
oxygen. Oxygen and carbon dioxide diffuse through their skin.
Many, not all, adult amphibians have gills for "breathing"
oxygen in the water. Often, the gills and skin are working
simultaneously as respiration organs.
Therefore, the skin has to be counted as an organ that in some
tetrapods is a respiratory organ. Some frogs have extremely
wrinkled skin to provide more surface area for the gas exchange.
Therefore, the skin in these cases is highly modified as a respiratory
organ.
Some turtles have highly vasuclarized tails. So oxygen is absorbed
in the water by the skin of their tails.

> Or do tadpoles qualify?
I think they do. Natural selection acts on all stages of development
of an animal. Therefore, one shouldn't restrict ones analysis only to the
adult stage. A feature can be useful at any stage of development, and
still be shaped by natural selection.

[jillery]

On Sat, 12 May 2012 14:30:56 -0400, Richard Norman

All very good examples of tetrapods with auxiliary water breathing
apparatus. Thank you for your reply.

[Paul Ciszek]

In article <20088799.879.1336857660465.JavaMail.geo-discussion-forums@ynja13>,

<drose...@yahoo.com> wrote:
> Adult amphibians generally "breath" through their skin. Their
>skin does not have any scales. The skin is moist and covered with
>oxygen. Oxygen and carbon dioxide diffuse through their skin.
> Many, not all, adult amphibians have gills for "breathing"
>oxygen in the water. Often, the gills and skin are working
>simultaneously as respiration organs.
> Therefore, the skin has to be counted as an organ that in some
>tetrapods is a respiratory organ. Some frogs have extremely
>wrinkled skin to provide more surface area for the gas exchange.
>Therefore, the skin in these cases is highly modified as a respiratory
>organ.
> Some turtles have highly vasuclarized tails. So oxygen is absorbed
>in the water by the skin of their tails.

Tails? Only is a euphemistic sense:

http://en.wikipedia.org/wiki/Cloaca#Cloacal_respiration

[Paul Ciszek]

In article <ltoqq7d8ffrm1phti...@4ax.com>,

Richard Norman <r_s_n...@comcast.net> wrote:
>
>You are quite correct in emphasizing the use of breathing organs other
>than lungs/swim bladders. I just mentioned it in passing and it is
>too easily passed by. I interpreted the question on obligate air
>breathing technically and narrowly without regard to what organ is
>involved. The real question is the relation of tetrapod lungs to fish
>lungs and swim bladders.

No, I asked the multiple questions that I did because I am interested
in air-breathing fish in general, including those who lost their lungs
and evolved other ways to breath air. Particularly interesting are
those cases of fish that *must* breath air, but have to do so using
something other than lungs.

--
Please reply to: | "We establish no religion in this country, we
pciszek at panix dot com | command no worship, we mandate no belief, nor
Autoreply is disabled | will we ever. Church and state are, and must
| remain, separate." --Ronald Reagan, 10/26/1984

[Richard Norman]

On Sun, 13 May 2012 14:10:56 +0000 (UTC), nos...@nospam.com (Paul
Ciszek) wrote:

>
>In article <ltoqq7d8ffrm1phti...@4ax.com>,
>Richard Norman <r_s_n...@comcast.net> wrote:
>>
>>You are quite correct in emphasizing the use of breathing organs other
>>than lungs/swim bladders. I just mentioned it in passing and it is
>>too easily passed by. I interpreted the question on obligate air
>>breathing technically and narrowly without regard to what organ is
>>involved. The real question is the relation of tetrapod lungs to fish
>>lungs and swim bladders.
>
>No, I asked the multiple questions that I did because I am interested
>in air-breathing fish in general, including those who lost their lungs
>and evolved other ways to breath air. Particularly interesting are
>those cases of fish that *must* breath air, but have to do so using
>something other than lungs.

I haven't gone back to the sources to verify this but my impression is
that pretty much all the respiratory exchange surfaces of fish
were/are derived from the pharynx or other regions of the digestive
tract There is good reason for that. The early ostracoderms and then
placoderms were heavily plated so that their external surface could
not be used as for respiratory exchange. By contrast, the pharyngeal
slits were originally used for both feeding and then for respiration
as gills. The mouth is also an easy way to take in air for surface
living fish. So the swim bladder and the fish lungs were both
pharyngeal derivations, as are most of the other breathing methods.
The buccal cavity and gut are also well vascularized surfaces that can
be used. The labyrinthine organ of gourami or Siamese fighting fish
is, I believe, a derivative of the gills. I have read that some eels
can use their skin for some respiration.

Fish are somewhat limited in that these respiratory surfaces have
limited area for exchange. Creating large areas for exchange is very
difficult because surface tension will collapse small air sacs. True
fish lungs have a surfactant coating to help prevent this much like
that of tetrapod lungs. The auxiliary respiratory exchange surfaces
in fish don't have surfactant and so cannot be finely divided and
still hold air.

[UC]

On May 13, 3:50 pm, Richard Norman <r_s_nor...@comcast.net> wrote:
> On Sun, 13 May 2012 14:10:56 +0000 (UTC), nos...@nospam.com (Paul
>
>
>
>
>
>
>
>
>
> Ciszek) wrote:
>

> >In article <ltoqq7d8ffrm1phtiper9j45eosp9d8...@4ax.com>,

I was not aware previously that there were any obligate air-breathing
fish.

[Richard Norman]

See, you can actually learn something here.

You can learn a lot more simply by learning to google, for example,
"obligate air breathing fish".

[pnyikos]

On May 11, 4:45 pm, Richard Norman <r_s_nor...@comcast.net> wrote:
> On Fri, 11 May 2012 15:35:31 -0400, jillery <69jpi...@gmail.com>

> wrote:
>
> >On Fri, 11 May 2012 12:57:38 -0400, Richard Norman
> ><r_s_nor...@comcast.net> wrote:
>
> <snip except relation between fish lungs and swim bladders>
> >> Is anything ever undisputed?  There are sufficient differences
> >>between the structure and development of fish lungs and fish swim
> >>bladders to put doubt on their relationship.  See, for example,
>
> >>   Which came first, the lung or the breath?
> >>  http://www.sciencedirect.com/science/article/pii/S109564330100304X
>
> >>The abstract says: "Lungs are paired ventral derivatives of the
> >>pharynx posterior to the gills.

Not in the Australian lungfish, see below.

I would count the following as evidence, however weak: the Australian
lungfish has a single dorsal (though still connected vertrally) lung
which I think is generally believed to be homologous to one of the
lungs of the other Dipnoi, of the bichir, and perhaps some other fish.

Query: is the swimbladder connected to the gut in most actinopterygian
fishes?

Peter Nyikos

>The most plausible hypothesis is that both have their origins
> in a “respiratory pharynx” (Neumayer 1930; Wassnetzov 1932). Thus, it
> is unlikely that lungs and the swimbladder are derived one from the
> other, since none of the three homology criteria (same relative
> location, same embryological origin, ontogenetic, or phylogenetic
> continuum) used in comparative anatomy and evolutionary biology are
> satisfied. Even the discovery of activity of the same genes in lung
> and swimbladder development would not prove the homology of the
> organs: hand and foot are not homologous although there genetic

> patterns of development are similar.- Hide quoted text -
>
> - Show quoted text -

[Richard Norman]

Why did you move this thread from talk.origins?

The lungfish belong to the Dipnoi. I am not sure whether that refers
to a double lung or double "breathing" (air and water). Still here is
a citation that indicates that Neoceratodus does have a single lung
whereas Lepidosiren and Protopterus have paired lungs. But
Neoceratodus has a vestige of a second lung as an embryo and the
single lung is considered a derived feature.

Circulation and Respiration in Lungfishes (Dipnoi)
W Burggren and K Johansen
J. Morphol., 190: 217–236 (1986)
doi: 10.1002/jmor.1051900415

http://www.cep.unt.edu/~burggren/pdfs/1986/(56)Burggren,Johansen1986JMS.pdf

In other words, the Australian lungfish, like so many things
Australian, is just plain weird. Neither the South American nor the
African lungfish are the same.

The phystostome swim bladder is connected to the gut (the pharynx is
part of the gut) while the physoclist swim bladder is separated. The
muskie (muskellunge) is a phystostome while perch and walleye are
physiclists. I don't know exactly what the relative distribution of
the two types of swim bladder is.

[Richard Norman]

I am copying this from sci.bio.paleontology because pnyikos posted it
there in addition to talk.origins but it doesn't seem to show up here.
My impression is that the sci.bio groups may be moderated and t.o.
doesn't allow cross posting to moderated groups. Then again, maybe it
will just show up here later in which case just ignore one of the
copies.

On Mon, 14 May 2012 16:54:21 -0700 (PDT), pnyikos
<nyi...@bellsouth.net> wrote:

>
>

>On May 11, 4:45 pm, Richard Norman <r_s_nor...@comcast.net> wrote:
>> On Fri, 11 May 2012 15:35:31 -0400, jillery <69jpi...@gmail.com>

>> wrote:
>>
>> >On Fri, 11 May 2012 12:57:38 -0400, Richard Norman
>> ><r_s_nor...@comcast.net> wrote:
>>
>> <snip except relation between fish lungs and swim bladders>
>> >> Is anything ever undisputed?  There are sufficient differences
>> >>between the structure and development of fish lungs and fish swim
>> >>bladders to put doubt on their relationship.  See, for example,
>>
>> >>   Which came first, the lung or the breath?
>> >>  http://www.sciencedirect.com/science/article/pii/S109564330100304X
>>
>> >>The abstract says: "Lungs are paired ventral derivatives of the
>> >>pharynx posterior to the gills.
>

>Not in the Australian lungfish, see below.
>
>

>I would count the following as evidence, however weak: the Australian
>lungfish has a single dorsal (though still connected vertrally) lung
>which I think is generally believed to be homologous to one of the
>lungs of the other Dipnoi, of the bichir, and perhaps some other fish.
>
>Query: is the swimbladder connected to the gut in most actinopterygian
>fishes?

Why did you move this thread from talk.origins?

The lungfish belong to the Dipnoi. I am not sure whether that refers
to a double lung or double "breathing" (air and water). Still here is
a citation that indicates that Neoceratodus does have a single lung
whereas Lepidosiren and Protopterus have paired lungs. But
Neoceratodus has a vestige of a second lung as an embryo and the
single lung is considered a derived feature.

Circulation and Respiration in Lungfishes (Dipnoi)
W Burggren and K Johansen
J. Morphol., 190: 217–236 (1986)
doi: 10.1002/jmor.1051900415

http://www.cep.unt.edu/~burggren/pdfs/1986/(56)Burggren,Johansen1986JMS.pdf

In other words, the Australian lungfish, like so many things
Australian, is just plain weird. Neither the South American nor the

African lungfish are the same; these have two lungs.

[Walter Bushell]

In article <bpn2r71sl8u7c9gh9...@4ax.com>,

Few would spontaneously thin of the concept "obligate air breathing
fish". Of course, Homer the sap is an > "obligate air breathing fish".

--
This space unintentionally left blank.

[Richard Norman]

On Tue, 15 May 2012 10:50:30 -0400, Walter Bushell <pr...@panix.com>
wrote:

If you really wanted to gather together all things called "fish" into
a single clade, then spiders and earthworms are obligate air breathing
fish.

[Arkalen]

If we really wanted to gather all things called "fish" into a single
clade, what organisms would NOT be fish ?
Plants ?

[Ernest Major]

In message <JWB*D+...@news.chiark.greenend.org.uk>, Arkalen
<ark...@inbox.com> writes

Sponges, fungi, algae, unicellular eukaryotes, bacteria, archaea

Now, if you wanted to gather all things called "slime moulds" into a
single clade ...

http://en.wikipedia.org/wiki/Slime_mould
--
Alias Ernest Major

[Richard Norman]

Starkist tuna is packed in a fish plant!

[Walter Bushell]

In article <JWB*D+...@news.chiark.greenend.org.uk>,

Insects, crabs, all the unsegmented forms, fungi and single cells, for
examples. Nearly all life forms.

[Richard Norman]

On Tue, 15 May 2012 21:51:41 -0400, Walter Bushell <pr...@panix.com>

Not if the clade includes cuttlefish, starfish, jellyfish, ...
That still leaves some animals plus all the other kingdoms.

[Don Cates]

What if you include fishweed? (B-)
<http://plantnet.rbgsyd.nsw.gov.au/cgi-bin/NSWfl.pl?page=nswfl&lvl=sp&name=Einadia~trigonos>

--
--
Don Cates ("he's a cunning rascal" PN)

[Ernest Major]

In message <proto-6706B4....@news.panix.com>, Walter Bushell
<pr...@panix.com> writes

>In article <JWB*D+...@news.chiark.greenend.org.uk>,
> Arkalen <ark...@inbox.com> wrote:
>
>> On 15/05/12 16:02, Richard Norman wrote:
>> > On Tue, 15 May 2012 10:50:30 -0400, Walter Bushell <pr...@panix.com>
>> > wrote:
>> >
>> >> In article <bpn2r71sl8u7c9gh9...@4ax.com>,
>> >> Richard Norman <r_s_n...@comcast.net> wrote:
>> >>
>> >>> On Mon, 14 May 2012 07:59:34 -0700 (PDT), UC
>> >>> <uraniumc...@yahoo.com> wrote:
>> >>>
>> >>>> On May 13, 3:50 pm, Richard Norman <r_s_nor...@comcast.net> wrote:
>> >>>>> On Sun, 13 May 2012 14:10:56 +0000 (UTC), nos...@nospam.com (Paul
>> >>>>>
>> >>>>>
>> >>>>> Ciszek) wrote:
>> >>>>>
>> >>>>>> In article <ltoqq7d8ffrm1phtiper9j45eosp9d8...@4ax.com>,
>> >>>>>> Richard Norman <r_s_nor...@comcast.net> wrote:
>> >>>>>
>> >>>>>>> You are quite correct in emphasizing the use of breathing
>> >>>>>>>organs other
>> >>>>>>> than lungs/swim bladders. I just mentioned it in passing and it is
>> >>>>>>> too easily passed by. I interpreted the question on obligate air
>> >>>>>>> breathing technically and narrowly without regard to what organ is
>> >>>>>>> involved. The real question is the relation of tetrapod

Cuttlefish would pull in insects and crabs and the other protostomes.
Jellyfish pulls in all animals other than sponges.
--
alias Ernest Major

[Richard Norman]

On Tue, 15 May 2012 22:59:06 -0500, Don Cates <caHOR...@mts.net>
wrote:

Excellent! My google skills were insufficient to dredge up that one.
Score another weirdness for those down under.

[drose...@yahoo.com]

On Monday, May 14, 2012 10:59:34 AM UTC-4, UC wrote:

> I was not aware previously that there were any obligate air-breathing
> fish.

There are a large number fresh water fish species that breath gaseous
air. There are a surprising number of fish species that are obligate
air breathers for all seasons.
I worked as a docent for an aquarium. I liked to point out the
air breathing fish to visitors. The most obvious air breather was "Sparky"
the electric eel. Sparky surfaced about once every ten minutes to take a
gulp of air. The fish near the surface swam to the other side of the tank
every time Sparky surfaced.
A lot of visitors were amazed at Sparky's air breathing abilities. I
was not able to demonstrate his electrical powers, but the air breathing
was very obvious. Like you, most visitors were not aware that there were
obligate air breathers among fish.

Here are three types of "obligate air-breathing fish" that
always need air.
1) Electric eels
2) Mud skipper
3) Snake head
These fish HAVE to surface occasionally, even if the water
is aerated. Note that the "mud skipper" spends 50-90% of its time
out of water. The mud skipper is not an amphibian. It is a
ray-finned fish that prefers to crawl out of the water most of
the time.

It should be noted that any fish that breathes air is ALWAYS an
obligate air breather in anoxic environments. Having the ability to
breath under water is not very useful when there is no oxygen in the
water. Water can become anoxic when the water becomes warm or when
there is a lot of decaying vegetation in the water.
Both the temperature of the water and the amount of dead vegetation
often varies with the time of year. In many bodies of fresh water, the water become anoxic during summer. Fish living in these waters have evolved
to breath air, hibernate or migrate. During the summer, any fish swimming
in these waters are obligate air breathers.
Fish that can both breath air and breath underwater include:
4) Carp (including goldfish)
5) Catfish (all of them)
6) Gar
7) Gourami

These fish prefer to stay underwater as long as the water is aerated.

[pnyikos]

On May 21, 6:12 pm, drosen0...@yahoo.com wrote:
> On Monday, May 14, 2012 10:59:34 AM UTC-4, UC wrote:
> > I was not aware previously that there were any obligate air-breathing
> > fish.
>
>      There are a large number fresh water fish species that breath gaseous
> air. There are a surprising number of fish species that are obligate
> air breathers for all seasons.
>      I worked as a docent for an aquarium. I liked to point out the
> air breathing fish to visitors. The most obvious air breather was "Sparky"
> the electric eel. Sparky surfaced about once every ten minutes to take a
> gulp of air. The fish near the surface swam to the other side of the tank
> every time Sparky surfaced.
>       A lot of visitors were amazed at Sparky's air breathing abilities. I
> was not able to demonstrate his electrical powers, but the air breathing
> was very obvious. Like you, most visitors were not aware that there were
> obligate air breathers among fish.
>
> Here are three types of "obligate air-breathing fish" that
> always need air.
> 1) Electric eels
> 2) Mud skipper
> 3) Snake head

The African lungfish is another.

>       These fish HAVE to surface occasionally, even if the water
> is aerated. Note that the "mud skipper" spends 50-90% of its time
> out of water. The mud skipper is not an amphibian. It is a
> ray-finned fish that prefers to crawl out of the water most of
> the time.
>
>      It should be noted that any fish that breathes air is ALWAYS an
> obligate air breather in anoxic environments. Having the ability to
> breath under water is not very useful when there is no oxygen in the
> water.  Water can become anoxic when the water becomes warm or when
> there is a lot of decaying vegetation in the water.
>       Both the temperature of the water and the amount of dead vegetation
> often varies with the time of year. In many bodies of fresh water, the water become anoxic during summer. Fish living in these waters have evolved
> to breath air, hibernate or migrate. During the summer, any fish swimming
> in these waters are obligate air breathers.
>      Fish that can both breath air and breath underwater include:
> 4) Carp (including goldfish)
> 5) Catfish (all of them)
> 6) Gar
> 7) Gourami

8) the Australian lungfish

Peter Nyikos

[pnyikos]

On May 16, 9:10 am, Richard Norman <r_s_nor...@comcast.net> wrote:
> On Tue, 15 May 2012 22:59:06 -0500, Don Cates <caHORMEL...@mts.net>

> wrote:
>
>
>
>
>
> >On 15/05/2012 9:42 PM, Richard Norman wrote:
> >> On Tue, 15 May 2012 21:51:41 -0400, Walter Bushell<pr...@panix.com>
> >> wrote:
>
> >>> In article<JWB*D+...@news.chiark.greenend.org.uk>,

> >>> Arkalen<arka...@inbox.com>  wrote:

>
> >>>> On 15/05/12 16:02, Richard Norman wrote:
> >>>>> On Tue, 15 May 2012 10:50:30 -0400, Walter Bushell<pr...@panix.com>
> >>>>> wrote:
>

> >>>>>> In article<bpn2r71sl8u7c9gh9grk65mt7t2rlbt...@4ax.com>,
> >>>>>> Richard Norman<r_s_nor...@comcast.net>  wrote:
>

> >>>>>>> On Mon, 14 May 2012 07:59:34 -0700 (PDT), UC

> >>>>>>> <uraniumcommit...@yahoo.com>  wrote:

> ><http://plantnet.rbgsyd.nsw.gov.au/cgi-bin/NSWfl.pl?page=nswfl&lvl=sp&...>

>
> Excellent!  My google skills were insufficient to dredge up that one.
> Score another weirdness for those down under.

Close, but no cigar. The name indicates that it is a weed, just as
spider crabs are crabs, while crab spiders are spiders.

Peter Nyikos

[pnyikos]

On May 14, 8:34 pm, Richard Norman <r_s_nor...@comcast.net> wrote:
> I am copying this from sci.bio.paleontology because pnyikos posted it
> there in addition to talk.origins but it doesn't seem to show up here.
> My impression is that the sci.bio groups may be moderated and t.o.
> doesn't allow cross posting to moderated groups.

It is sci.bio.evolution that does not allow crossposting to
talk.origins; sci.bio.paleontology is unmoderated, as a glance at its
table of contents will show. I've even restored the newsgroup.

>Then again, maybe it
> will just show up here later in which case just ignore one of the
> copies.

Well, I certainly won't re-post it here.

I can't recall whether I did or not. I probably crossposted it, then
part got confined to s.b.p.

> The lungfish belong to the Dipnoi. I am not sure whether that refers
> to a double lung or double "breathing" (air and water).  Still here is
> a citation that indicates that Neoceratodus does have a single lung
> whereas Lepidosiren and Protopterus have paired lungs.  But
> Neoceratodus has a vestige of a second lung as an embryo and the
> single lung is considered a derived feature.

All of which tends to support the hypothesis that the single swim
bladder did evolve from one of a set of paired lungs.

>   Circulation and Respiration in Lungfishes (Dipnoi)
>  W  Burggren and K Johansen
>   J. Morphol., 190: 217–236 (1986)
>   doi: 10.1002/jmor.1051900415
> http://www.cep.unt.edu/~burggren/pdfs/1986/(56)Burggren,Johansen1986JMS.pdf

Thank you for the reference. Dipnoi have been one of my favorite
subjects since I saw a live Australian lungfish at the San Francisco
aquarium when I was twelve years old. That experience ranks right up
with my first visit to the American Museum of Natural History later
that year.

> >
> In other words, the Australian lungfish, like so many things
> Australian, is just plain weird.  Neither the South American nor the
> African lungfish are the same; these have two lungs.
>
> The phystostome swim bladder is connected to the gut (the pharynx is
> part of the gut) while the physoclist swim bladder is separated.  The
> muskie (muskellunge) is a phystostome while perch and walleye are
> physiclists.  I don't know exactly what the relative distribution of
> the two types of swim bladder is.

Nor I, but it's something I am very curious about. If you come across
a good reference, let me know.

Peter Nyikos

[Richard Norman]

Everything spins backwards down under so the rule of which noun is
dominant gets reversed.

[Richard Norman]

>> The lungfish belong to the Dipnoi. I am not sure whether that refers
>> to a double lung or double "breathing" (air and water).  Still here is
>> a citation that indicates that Neoceratodus does have a single lung
>> whereas Lepidosiren and Protopterus have paired lungs.  But
>> Neoceratodus has a vestige of a second lung as an embryo and the
>> single lung is considered a derived feature.
>
>All of which tends to support the hypothesis that the single swim
>bladder did evolve from one of a set of paired lungs.

But I cited a tremendous amount of material which suggests
differently. Just because one kind of lungfish lost its second lung
doesn't mean that the single swim bladder is also a relic of one of
the presumably paired lungs of an early fish-like thing.

[Richard Norman]

Phystostomous fish, with the swim bladder connected to the gut,
swallow air so they obvious are fish that come up to the surface.
Physoclistous fish, on the other hand, have to secrete and absorb
gases from and into the blood to control their swim bladder and so
cannot do that rapidly. Therefore they cannot change swim depth
quickly. Many are deep water animals that die if pulled up to the
surface too quickly.

Examples of phystostomous: catfish, trout and salmon, sturgeon,
minnows, suckers, and eels.

Examples of phystoclitous: bass, perch, mosquitofish, and sunfish,
rockfish.

Taken from http://chamisa.freeshell.org/fish.htm

I cannot find any source with a complete list of the two types or
whether there is any taxonomic division. If you search out the
families of all the examples I have given you, you might find some
division. I don't have the patience (or the interest) to do all that
work.

[funfis...@gmail.com]

http://www.youtube.com/watch?v=361xRfAO5W8