Proposed Sympatry of Midas and Red Devil Cichlids

New cichlid species and taxonomy
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Proposed Sympatry of Midas and Red Devil Cichlids

Post by RobertPrice » Mon Aug 06, 2012 12:49 pm

In this article, I propose to use modern sytematic theory that was not available to Gunther (1864) to give credible evidence that the Midas/Red Devil Complex in Lake Nicaragua is nothing more than two extreme races of the same species. I use the word races deliberately although not commonly used in taxonomy except for human beings, as the situation in the Crater Lakes of Nicaragua is exceptional and races will make my proposal easier to digest.

Red Devils Amphilophus labiatusand Midas cichlids Amphilophus citrinellus were both described by Gunther in 1864. Gunther was hampered by having only a dearth of museum specimens, a lack of knowledge of the complex phylogeography of these fish, and because he was in the pre-DNA era, unable to use this most valuable indicator of speciation. These species' genetics were extensively studied by Barluenga and Meyer (2010) who elucidated several new species of the complex in nearby minor crater lakes, yet after having stated that [citrinellusand and labiatus DNA were virtually indistinguishable in Lake Nicaragua, did not emphasize that the two freely interbreed in the lake, in which their ancestral form has been for at least 100,000 years. Seth Meek (1907), stated that despite his efforts, he could not recognize two distinct species of this complex in Lake Nicaragua. Add to this Barleunga and Meyer's (2010) statements, by recognizing twp taxonomic entities you fly in the face of the systematists guideline of parsimony: that the simplest explanation is likely the most correct.

In the online article Midas Cichlid, Amphilophus citrinellus Animal-World speculates the Red Devils and Midas are the the same species, because the majority of both are sympatric in Lake Nicaragua and look and act very much alike. (Midas are found in Lake Managua, the small Nicarguan crater Lakes and Costa Rica as well, and Red Devils, though not many, are found in Managua). They correctly note that both are avid diggers on the bottom, and Barleunga and Meyer (2010) admit that "few credible examples of sympatric speciation exist," and that Midas and Red Devils "remain genetically surprisingly indistinguishable."

When encountered, geophraphic isolation and the inability to interbreed define two species. Genetic drift within similar organisms that share most of their geographic range requires a more philosphic approach. As yet supposed preferences about living at different heights in the water column are not terribly significant. If I live on Bear Mountain and my friends live in the Bronx, does that indicate we are allopatric or less closely related? Of course not. The argument about Red Devil's having thicker lips so are a distinct species is poppycock. Within the human species we have obvious differences in lip thickness in breeding populations. it has been stated that most Midas cichlids are higher or taller than most Red Devils. Several African tribes of humans average a foot taller than some Oriental people. Another specious argument is the size of the hump. Let's not play "mine is bigger than yours.' that is for children. Many species of male cichlids have humps, and there are usually clines within a species ranging from the biggest to smallest humps. Finally, both species are polychromatic and come in lots of shades of several colors. So do people, and I don't know of any scientific proposal that there are several human species.

In short, there are as many differences in races of [Homo sapiens as there are between citrinellus and labiatus Just as all people can and often do breed, so do Red Devils and Midas Cichlids Barleugena (2006, 2010) and his associates have stated that Nicaraguan Crater Lake cichlids of the Midas Complex are a great model for incipient speciation and defining a species flock. I have no problems wiith zaliuosus and the other nearby geographically isolated species they describe, but as long as Lake Nicaragua survives,citrinellus and labiatus are in the same position as different human races who are mobile within each other's areas of greatest concentration, and freely breed. It is indeed curious that Barluenga et al, after their exhaustive DNA analysis, have to fall back on small morphological differences to hustify separation of Midas and Red Devil cichlids.

Albert Gunther, the greatest ichthyologist of his age, described both fish in 1864, but designated the type speicmens of Red Devils, whicfh he named labiatum in 1863. Therefore, citrinellusbecomes a junior synonym of labiatus, and we now designate the Nicaraguan speciesAmphilophus labiatus as monotyptic and composed of two races, labiatus and citrinellus.

References:

M. Barleunga, N. Stotting, W. Salzburger M Mishnick and A. Meyer. 2006. Sympatric speciation in Nacaraguan crater lake cichlids. Nature, Feb 9, (7077)19-23

Marta Barleunga and A. Meyer. Phylogeography, colonization, and population history of the Midas Cichlid Species Complex Amphilophus spp. in Nacaraguan crater lakes. Evolutionary Biology 2010 10:326

Meek, Seth 1907. Synopsis of the fishes of the great lakes of Nicaragua. Field Columbian Museum zoological series 1907, publ. 121 7:97-121

Midas Cichlid, Amphilophus citrinellus Animal-World
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Re: Proposed Sympatry of Midas and Red Devil Cichlids

Post by RobertPrice » Tue Aug 07, 2012 11:16 am

I'd like to correct my title. It's "Proposed Synonomy" not Sympatry. Me bad. Honnestly my eyesight is not the best and I can't read my own handwritiing. :?

Dr. Robert Price, Ph.d,, Herpetology and Ichthyology
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Re: Proposed Sympatry of Midas and Red Devil Cichlids

Post by Willem Heijns » Tue Aug 07, 2012 1:52 pm

Dear Robert,

I have read your contribution a couple of times, but I can't seem to find the "credible evidence" for synonymy of Amphilophus citrinellus and Amphilophus labiatus in it.
Many if not most of the things you say apply to all species of the so-called citrinellus complex, whether they occur in the Great Lakes of Nicaragua or in one or more of the smaller crater lakes (where sympatric spciation has most likely occurred on several occasions).

Would you be so kind as to summarize the characters on which you have based your opinion?
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Re: Proposed Sympatry of Midas and Red Devil Cichlids

Post by gnuisance » Tue Aug 07, 2012 2:21 pm

I enjoyed reading your article and would not challenge your findings. My only feedback would be that you get away from your "scientific voice" in the fourth paragraph. The casual language and slang that you use takes away a bit from the credibility of the paper. Other than that I think it was very well done.

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Re: Proposed Sympatry of Midas and Red Devil Cichlids

Post by Bas Pels » Tue Aug 07, 2012 2:29 pm

In all honesty, I can not see any argument in the first posting which convinces me

Amphilophus citrinellum and A. labiatum can interbreed. So what? I've seen Hypsophrys nicaraquensae and Cryptoheros nigrocfasciatus interbreed. Amphilophus trimaculatus ans "Cichlasoma istlanum" interbred in my house - and a year later the offspring also produced swimming fry. I found a hybrid of Herichthys labiatum (although, it could have been H panosticticus) with Cryptoheros septemfasciatus in a swarm of C. septemfasciatus. This is not an argument.

All cichlids of Central america are suspected of being able to interbreed. Species have to be defined differently. And this possible interbreeding might even be the reason why it is so hard to draw conclusions from DNA searches: they quite often contradict each other, perhaps due to this possibility of dealing with hybrids, which later crossed back into one of the parent species. I could not say.

If one would compare a man of the pygmy tribes with a man from Norway, I'm quite certain a extra terrestrial taxonimist, who would know nothing about humans, would from a morphological point of view think about two species - but we know better. Further, all taxonimical research in the genus Homo has to do with sensitive politics, and therefore I think it is better to keep Homo sapiens out of any taxonomical discussion.

I do agree A citrinellum and A. labiatum are very closely related, and they might even be the same species - but that would require better arguments

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Re: Proposed Sympatry of Midas and Red Devil Cichlids

Post by RobertPrice » Tue Aug 07, 2012 2:59 pm

Willem Heijns wrote:Dear Robert,

I have read your contribution a couple of times, but I can't seem to find the "credible evidence" for synonymy of Amphilophus citrinellus and Amphilophus labiatus in it.
Many if not most of the things you say apply to all species of the so-called citrinellus complex, whether they occur in the Great Lakes of Nicaragua or in one or more of the smaller crater lakes (where sympatric spciation has most likely occurred on several occasions).

Would you be so kind as to summarize the characters on which you have based your opinion?
The characters on which I based my conclusion are Barleunga's findings that Lake Nicaragua Red Devils amd Midas cichlids have essentially the same DNA, dont look dissimilar enough to justify speciation, and constantly interbrred. The accepted definition of species is that they do not breed except possibly at the convergence of their geographic ranges, share common characteristics, and are sexually isolated. the definition of subspecies provides for separate geographic ranges. LThe vast majority of Nicaraguan Midas and Red Devils do no't have separate geographic ranges, do interbreed, and have essentially the same DNA. if you Google "Images of Midas Cichlids," and "Images of Red Devils," you will see that you are looking at variations of the same fish. The small crater like species such as Zaliousus are Midas cichlid offshoots, but they no longer interbreed with Midas, and most are vertically heavily striped and look more like big Convict cichlids.
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Re: Proposed Sympatry of Midas and Red Devil Cichlids

Post by RobertPrice » Tue Aug 07, 2012 3:03 pm

gnuisance wrote:I enjoyed reading your article and would not challenge your findings. My only feedback would be that you get away from your "scientific voice" in the fourth paragraph. The casual language and slang that you use takes away a bit from the credibility of the paper. Other than that I think it was very well done.
I admit I did it deliberately to make my point more easy to understand to people who are not very familiar with systematics and taxonomy, and the political correctness that is applied to humans, but often ignored with other animals. Thank you for your comments and critique.
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Re: Proposed Sympatry of Midas and Red Devil Cichlids

Post by RobertPrice » Tue Aug 07, 2012 3:24 pm

Bas Pels wrote:In all honesty, I can not see any argument in the first posting which convinces me

Amphilophus citrinellum and A. labiatum can interbreed. So what? I've seen Hypsophrys nicaraquensae and Cryptoheros nigrocfasciatus interbreed. Amphilophus trimaculatus ans "Cichlasoma istlanum" interbred in my house - and a year later the offspring also produced swimming fry. I found a hybrid of Herichthys labiatum (although, it could have been H panosticticus) with Cryptoheros septemfasciatus in a swarm of C. septemfasciatus. This is not an argument.

All cichlids of Central america are suspected of being able to interbreed. Species have to be defined differently. And this possible interbreeding might even be the reason why it is so hard to draw conclusions from DNA searches: they quite often contradict each other, perhaps due to this possibility of dealing with hybrids, which later crossed back into one of the parent species. I could not say.

If one would compare a man of the pygmy tribes with a man from Norway, I'm quite certain a extra terrestrial taxonimist, who would know nothing about humans, would from a morphological point of view think about two species - but we know better. Further, all taxonimical research in the genus Homo has to do with sensitive politics, and therefore I think it is better to keep Homo sapiens out of any taxonomical discussion.

I do agree A citrinellum and A. labiatum are very closely related, and they might even be the same species - but that would require better arguments
Barluenga et al. spent 4 years testing the DNA of Red Devils and Midas Cichlids and concluded that they are almost exactly the same. The genetic differences in humans that account for presence or absence of an Epicanthic Fold over the eye are as great as has been found between these two fish. Lake Nicaragua is a big like and the Midas cichlids in different parts of the Lake look different. so do the Red Devils, but they share the territory of the entire lake and interbrred all the time. None of the other Central American cichlids do that.

After 40 years of experience as a professional lower vertebrate systematist, i've been dismayed by colleagues who are constantly separating and subspeciating everything based on puerile reasoning just to get their names in print. Seth Meek was a great ichthyologist, and he said they were the same, but never bothered to describe the synonomy. I respect him for this as he and Gunther were working with museum specimens. I've done the same with kingsnakes of the species Lampropletis calligaster, and in 4 years found only two which were geographically and morphologically distinct. I described them as a subspecies. Perhaps citrinellus andlabiatus are slowly on their way to speciation, but they do not meet the criteria yet.
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Re: Proposed Sympatry of Midas and Red Devil Cichlids

Post by Nuchal Man » Tue Aug 07, 2012 11:46 pm

I to don't find credible evidence for synonymy and I don't think you make a very good argument. I believe you're also missing a relevant paper on the genus that was published in Molecular Phylogenetics and Evolution by Geiger et al., 2010. They found that "Lake Nicaragua’s species assemblage does not constitute a monophyletic group as it contains the four most basal taxa of the in-group, belonging to four different putative species". They also state that "treating all identified phenotypes and described species as members of a highly polymorphic A. citrinellus does also not reflect the discovered pattern and should be discarded". To not consider this paper while making this argument, even though this paper is discussed in Barleunga et. al., 2010 is a mistake. I really think what hinders many of the phylogenetic studies on cichlids is that they most use mitochondrial DNA. Many contradict each other as Bas stated. We know mitochondrial DNA is not the best in elucidating relationships between cichlids as they are rapidly evolving, hybridization is fairly common in the wild, and many are rather young radiations (including the midas cichlids). We also know that the analysis of mitochondrial DNA and Nuclear DNA can also yield quite different evolutionary processes when used to analyze the same group of fish!

There also is a statement in Barleunga et al., 2010 which seems to contradict your argument even though it is the main paper of your support. They state "in the laboratory thick-lipped fish prefer to mate with thick-lipped fish over fish with small lips (A.M. pers. obs.) further supporting their status as two described species".

Another point you bring up that is invalid is that "small crater like species such as Zaliousus are Midas cichlid offshoots, but they no longer interbreed with Midas, and most are vertically heavily striped and look more like big Convict cichlids". Amphilophus labiatus and Amphilophus citrinellus also have heavy baring. The midas cichlid complex is a highly color polymoprhic complex. Many of the small crater lake species also have been documented with color polymorphisms, for instance Amphilophus saggitae, Amphilophus amarillo, and Amphilophus xiloaensis (McKaye et al., 2002 & Geiger et al., 2010).

Your comments on hybridization are also intriguing. So what if they interbreed? Fish are going to hybridize every so often naturally no matter what, it is bound to happen, no matter how large the lake and no matter how many individuals of the same species there are. Many cichlids, not just Central American cichlids will hybridize with each other. Bas's observations of a variety of hybridization in captivity is quite true. Another great example that I'm sure he's probably seen but didn't mention is Paratheraps and Vieja. Hybrids between fish of these genera are quite common in captivity, not just intrageneric, but intergeneric hybrids as well. I'm sure Willem could also add his share of examples.

In the wild, we also see lots of hybridization in cichlids and it's been thought by quite a number of authors that this has helped provide a launching point for rapid speciation! Lake Tanganyikas Lamprologines (Sturmbauer et al., 2010 & Day et al., 2007) and Tropheini (Koblmüller et al., 2010 & Egger et al., 2007) are a fantastic examples. It's also been witnessed and video recorded in the wild, specifically by Heinz Buscher with Lamprologine cichlids. Lake Victoria we know has huge amounts of hybridization due to the deterioration of the lake and hybridization more than likely played a role before the lake began to deteriorate. Lake Malawi is another fantastic example of intergeneric hybridization in the wild (Genner & Turner, 2012). A colleague in an unpublished paper has also found that Astatotilapia calliptera pops up numerous in different parts of the Malawi cichlid phylogeny suggesting that there have been multiple times where Astatotilapia calliptera has entered the lake and hybridized with the endemic species. By your definition Astatotilapia and many of the mbuna and malawi Haplochromines would be considered the same species.


Literature Cited:

-Barleunga, M., N. Stotting, W. Salzburger M Mishnick and A. Meyer (2006) Sympatric speciation in Nacaraguan crater lake cichlids. Nature, 7077, 19-23.

-Day, J.J., Santini, S., Garcia-Moreno, J., (2007) Phylogenetic relationships of the Lake Tanganyika cichlid tribe Lamprologini: the story from mitochondrial DNA. Mol. Phylogenet. Evol. 46, 629-642.

-Egger, B., Koblmüller, S., Sturmbauer, C., Sefc, K.M., (2007) Nuclear and mitochondrial data reveal different evolutionary processes in the Lake Tanganyika cichlid genus Tropheus. BMC Evol. Biol. 7, 137.

- Geiger MF, McCrary JK, Schliewen UK (2010) Not a simple case - A first comprehensive phylogenetic hypothesis for the Midas cichlid complex in Nicaragua (Teleostei: Cichlidae Amphilophus). Molecular Phylogenetics and Evolution, 56,1011-1024.

-Genner, M.J. & G.F. Turner (2012) Ancient hybridisation and phenotypic novelty within Lake Malawi’s cichlid fish radiation. Molecular Biology and Evolution 29, 195–206.

- Kobmuller, s., Egger, B., Sturmbauer, C., Sefc, K.M. (2010) Rapid radiation, ancient incomplete lineage sorting and ancient hybridization in the endemic Lake Tanganyika cichlid tribe Tropheini. Molecular Phylogenetics and Evolution, 55, 318–334.

-McKaye, K.R., Stauffer Jr., J.R., van den Berghe, E.P., Vivas, R., Lopez Perez, L.J., McCrary, J.K., Waid, R., Konings, A., Lee, W.-J., Kocher, T.D., (2002) Behavioral, morphological and genetic evidence of divergence of the Midas cichlid species complex in two Nicaraguan crater lakes. Cuadernos de Investigación de la UCA 12, 19–47.

-Sturmbauer, C., Salzburger, W., Duftner, N., Schelly, R., Koblmüller, S.(2010) Evolutionary history of the Lake Tanganyika cichlid tribe Lamprologini (Teleostei: Perciformes) derived from mitochondrial and nuclear DNA data, Molecular Phylogenetics and Evolution, 57, 266-284.

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Re: Proposed Sympatry of Midas and Red Devil Cichlids

Post by Willem Heijns » Wed Aug 08, 2012 4:15 am

RobertPrice wrote: The characters on which I based my conclusion are Barleunga's findings that Lake Nicaragua Red Devils amd Midas cichlids have essentially the same DNA, dont look dissimilar enough to justify speciation, and constantly interbrred. The accepted definition of species is that they do not breed except possibly at the convergence of their geographic ranges, share common characteristics, and are sexually isolated. the definition of subspecies provides for separate geographic ranges. LThe vast majority of Nicaraguan Midas and Red Devils do no't have separate geographic ranges, do interbreed, and have essentially the same DNA. if you Google "Images of Midas Cichlids," and "Images of Red Devils," you will see that you are looking at variations of the same fish. The small crater like species such as Zaliousus are Midas cichlid offshoots, but they no longer interbreed with Midas, and most are vertically heavily striped and look more like big Convict cichlids.
Sorry Robert, your arguments are neither very convincing nor credible. You have made selective reference to the work of Barluenga (note the correct spelling) and Meyer. These authors firmly believe citrinellus and labiatus are separate species (albeit young species) as they state: "the extent of the genetic (my boldface) differences found here supports the assignment of Amphilophus citrinellus and Amphilophus labiatus as distinct biological species" (Barluenga & Meyer 2010: page 2071). Your statement "they don't look dissimilar enough" can hardly be considered scientific. Further, when you say: "they constantly interbreed" you are clearly implying field observations. I don't know if you have ever visited Lago Nicaragua and/or Lago Managua, but field observations there are impossible due to the turbidity of the water. Your advice to "google images of Midas cichlids" only brought a smile to my face. Is this how science is done? In conclusion, I don't think you have ever been to Laguna Apoyo either. Amphilophus zaliosus (again note the correct spelling) shows exactly the same (variation in) coloration as the other species of the citrinellus-complex in this lake do. In fact it almost never shows vertical barring. In normal coloration it only shows the body spots (an interrupted horizontal stripe as in all Amphilophus species) and in breeding coloration it is almost completely black. And although I must admit I have never seen zaliosus interbreed with other species in Laguna Apoyo on any of my dives there, the specimens that I took home with me readily interbred with an unrelated Amphilophus species from Laguna Monte Galán (a swamp north of Lago Managua) in the aquarium, even given the choice of a conspecific mate.
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Re: Proposed Sympatry of Midas and Red Devil Cichlids

Post by RobertPrice » Wed Aug 08, 2012 9:23 am

Willem Heijns wrote:Dear Robert,

I have read your contribution a couple of times, but I can't seem to find the "credible evidence" for synonymy of Amphilophus citrinellus and Amphilophus labiatus in it.
Many if not most of the things you say apply to all species of the so-called citrinellus complex, whether they occur in the Great Lakes of Nicaragua or in one or more of the smaller crater lakes (where sympatric spciation has most likely occurred on several occasions).

Would you be so kind as to summarize the characters on which you have based your opinion?
As far as your arguments about the small crater lakes go, they are true. citrinellus can be found with zaliosus the more recently described forms there. Mata Barleunga has stated that they do not socialize or interbreed, although they may be only 10,000 year old species. In the smaller lakes, the offshoots of Midas differentiate. Also, Midas in Lake Managua are genetically further from Midas in Lake Nicaragua than the L. Nicaragua natives are to Red Devils. There is only one tiny outlet from Lake Nicaragua, and in the fairly recent past it may have been larger and allowed peripherally dwelling Midas to migrate to the smaller Crater Lakes. The things I said about the relationship of Midas and Red Devils to not apply to the other forms of the complex. A few are sympatric with small colonies of Midas, and might breed in captivity, but they are so obviously different in appearance that few would be tempted to call them the same, and they do not interbreed except in fish tanks, which cannot be applied to a natural environmental scenario.

Finally if Marta Barleunga agrees that the two are "virtually indistnguishable." by regular and mitochondrial DNA anaylsis, she has as much as said they are the same species. She seems to be wanting to say so, but unwilling to rock Gunther's boat. Seth Meek, who disovered the Firemouth, was not nearly as overcautious, having said they are probably the same species now, but may diverge a good ways in the future, with which I agree.
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Re: Proposed Sympatry of Midas and Red Devil Cichlids

Post by Willem Heijns » Wed Aug 08, 2012 9:40 am

Robert,

Have you seen my second post? Or have you chosen to ignore it?
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Re: Proposed Sympatry of Midas and Red Devil Cichlids

Post by RobertPrice » Wed Aug 08, 2012 10:05 am

Willem Heijns wrote:Dear Robert,

I have read your contribution a couple of times, but I can't seem to find the "credible evidence" for synonymy of Amphilophus citrinellus and Amphilophus labiatus in it.
Many if not most of the things you say apply to all species of the so-called citrinellus complex, whether they occur in the Great Lakes of Nicaragua or in one or more of the smaller crater lakes (where sympatric spciation has most likely occurred on several occasions).

Would you be so kind as to summarize the characters on which you have based your opinion?
Sympatric speciation has by all accounts taken place in the smaller, much younger crater lakes in as little as 10,000 years. However, Midas and Red Devils have an ancestor in the large crater lakes as long as 100,000 years ago. If in this amount of time they have not done what the offshoot species did in 1/10 the time, there must be a reason. That reason is that there is enough gene flow back and forth between the two to preclude them from becoming distinct species. This was the reason for my analogy with humans. As far as sociologists can determine, Whites, Blacks and Orientals have a preference for breeding within their race rather than out-breeding, but they mix enough and do so constantly so that no one questions they are the same species despite minor morphological differences.
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Re: Proposed Sympatry of Midas and Red Devil Cichlids

Post by Willem Heijns » Wed Aug 08, 2012 10:14 am

It's hard to discuss something with someone who doesn't listen. So I'm out of this one.
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Re: Proposed Sympatry of Midas and Red Devil Cichlids

Post by RobertPrice » Wed Aug 08, 2012 11:25 am

Willem Heijns wrote:
RobertPrice wrote: The characters on which I based my conclusion are Barleunga's findings that Lake Nicaragua Red Devils amd Midas cichlids have essentially the same DNA, dont look dissimilar enough to justify speciation, and constantly interbrred. The accepted definition of species is that they do not breed except possibly at the convergence of their geographic ranges, share common characteristics, and are sexually isolated. the definition of subspecies provides for separate geographic ranges. LThe vast majority of Nicaraguan Midas and Red Devils do no't have separate geographic ranges, do interbreed, and have essentially the same DNA. if you Google "Images of Midas Cichlids," and "Images of Red Devils," you will see that you are looking at variations of the same fish. The small crater like species such as Zaliousus are Midas cichlid offshoots, but they no longer interbreed with Midas, and most are vertically heavily striped and look more like big Convict cichlids.
Sorry Robert, your arguments are neither very convincing nor credible. You have made selective reference to the work of Barluenga (note the correct spelling) and Meyer. These authors firmly believe citrinellus and labiatus are separate species (albeit young species) as they state: "the extent of the genetic (my boldface) differences found here supports the assignment of Amphilophus citrinellus and Amphilophus labiatus as distinct biological species" (Barluenga & Meyer 2010: page 2071). Your statement "they don't look dissimilar enough" can hardly be considered scientific. Further, when you say: "they constantly interbreed" you are clearly implying field observations. I don't know if you have ever visited Lago Nicaragua and/or Lago Managua, but field observations there are impossible due to the turbidity of the water. Your advice to "google images of Midas cichlids" only brought a smile to my face. Is this how science is done? In conclusion, I don't think you have ever been to Laguna Apoyo either. Amphilophus zaliosus (again note the correct spelling) shows exactly the same (variation in) coloration as the other species of the citrinellus-complex in this lake do. In fact it almost never shows vertical barring. In normal coloration it only shows the body spots (an interrupted horizontal stripe as in all Amphilophus species) and in breeding coloration it is almost completely black. And although I must admit I have never seen zaliosus interbreed with other species in Laguna Apoyo on any of my dives there, the specimens that I took home with me readily interbred with an unrelated Amphilophus species from Laguna Monte Galán (a swamp north of Lago Managua) in the aquarium, even given the choice of a conspecific mate.
I apoligize for my poor spelling, as my eyesight is poor. Your defense of your position is rife with self condictions and misconceptions. I am a Master Scuba
Diver and appreicate your trips to observe the Lakes, However, you yourself said Apoyo is too turbid to see anything important, then admit you have never seen zaliousus breeding there, but they breed with other fish at home. At home doesn't count. It is a perverion of the natural state that induces fish to behave out of character. Neither does the polytypy of the remaning 15 species of cichlids in the lake count. My paper only deals with Midas and Red Devils, who do not interbreed with the others as far as is known in the wild. I do not deny there has been siginificant speciation of other cichlids in the approximately half
million years since the prototypical cichlids arrived there.

You seem expert on the fish themselves, but lacking in an education in Systematics. Prior to the advent of DNA, the definition of a species was sympatric organisms that share most the same physical traits that breed amongst themselves except on the perihpery of their range with an adjoinig species. Similar looking sympatric organisms that interbreed and have indistnguishable DNA furthers the certainty that they are species. Races, mostly confined in sensu stricto to Humans, are not quite so similar looking members of the same species that have areas of sympatry and interbreed. Unless you care to appeal to International Commission on Zoological Nomenclature to change the definition of species, the vast preponderance of the evidence makes Midas and Red Devils races, not species.
Robert Price, Ph.d

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Re: Proposed Sympatry of Midas and Red Devil Cichlids

Post by RobertPrice » Wed Aug 08, 2012 1:26 pm

Willem Heijns wrote:It's hard to discuss something with someone who doesn't listen. So I'm out of this one.
Good Riddance!
Robert Price, Ph.d

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Re: Proposed Sympatry of Midas and Red Devil Cichlids

Post by Lisachromis » Wed Aug 08, 2012 3:11 pm

RobertPrice wrote:
Willem Heijns wrote:It's hard to discuss something with someone who doesn't listen. So I'm out of this one.
Good Riddance!
Please keep to the discussion. We prefer to keep things civil. I am moving the post into the taxonomy section where more people who have an interest in systemics will see the topic.

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Re: Proposed Sympatry of Midas and Red Devil Cichlids

Post by smitty » Wed Aug 08, 2012 8:21 pm

What an awesome topic that I know i will read at least 100's of times
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180gal- Managuense; 3 AC 110 P.Filters; 2 AC 110 P.Heads; Eheim 2217,2260; FX5
210gal- Argta,Bifsct,Fnstrtus,Maculcda,Regani,Syns,Zntus; 4 AC P.Filters; 2 AC P.Hds; Eheim 2250,2262; FX5

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Re: Proposed Sympatry of Midas and Red Devil Cichlids

Post by RobertPrice » Thu Aug 09, 2012 1:52 am

Nuchal Man wrote:I to don't find credible evidence for synonymy and I don't think you make a very good argument. I believe you're also missing a relevant paper on the genus that was published in Molecular Phylogenetics and Evolution by Geiger et al., 2010. They found that "Lake Nicaragua’s species assemblage does not constitute a monophyletic group as it contains the four most basal taxa of the in-group, belonging to four different putative species". They also state that "treating all identified phenotypes and described species as members of a highly polymorphic A. citrinellus does also not reflect the discovered pattern and should be discarded". To not consider this paper while making this argument, even though this paper is discussed in Barleunga et. al., 2010 is a mistake. I really think what hinders many of the phylogenetic studies on cichlids is that they most use mitochondrial DNA. Many contradict each other as Bas stated. We know mitochondrial DNA is not the best in elucidating relationships between cichlids as they are rapidly evolving, hybridization is fairly common in the wild, and many are rather young radiations (including the midas cichlids). We also know that the analysis of mitochondrial DNA and Nuclear DNA can also yield quite different evolutionary processes when used to analyze the same group of fish!

There also is a statement in Barleunga et al., 2010 which seems to contradict your argument even though it is the main paper of your support. They state "in the laboratory thick-lipped fish prefer to mate with thick-lipped fish over fish with small lips (A.M. pers. obs.) further supporting their status as two described species".

Another point you bring up that is invalid is that "small crater like species such as Zaliousus are Midas cichlid offshoots, but they no longer interbreed with Midas, and most are vertically heavily striped and look more like big Convict cichlids". Amphilophus labiatus and Amphilophus citrinellus also have heavy baring. The midas cichlid complex is a highly color polymoprhic complex. Many of the small crater lake species also have been documented with color polymorphisms, for instance Amphilophus saggitae, Amphilophus amarillo, and Amphilophus xiloaensis (McKaye et al., 2002 & Geiger et al., 2010).

Your comments on hybridization are also intriguing. So what if they interbreed? Fish are going to hybridize every so often naturally no matter what, it is bound to happen, no matter how large the lake and no matter how many individuals of the same species there are. Many cichlids, not just Central American cichlids will hybridize with each other. Bas's observations of a variety of hybridization in captivity is quite true. Another great example that I'm sure he's probably seen but didn't mention is Paratheraps and Vieja. Hybrids between fish of these genera are quite common in captivity, not just intrageneric, but intergeneric hybrids as well. I'm sure Willem could also add his share of examples.

In the wild, we also see lots of hybridization in cichlids and it's been thought by quite a number of authors that this has helped provide a launching point for rapid speciation! Lake Tanganyikas Lamprologines (Sturmbauer et al., 2010 & Day et al., 2007) and Tropheini (Koblmüller et al., 2010 & Egger et al., 2007) are a fantastic examples. It's also been witnessed and video recorded in the wild, specifically by Heinz Buscher with Lamprologine cichlids. Lake Victoria we know has huge amounts of hybridization due to the deterioration of the lake and hybridization more than likely played a role before the lake began to deteriorate. Lake Malawi is another fantastic example of intergeneric hybridization in the wild (Genner & Turner, 2012). A colleague in an unpublished paper has also found that Astatotilapia calliptera pops up numerous in different parts of the Malawi cichlid phylogeny suggesting that there have been multiple times where Astatotilapia calliptera has entered the lake and hybridized with the endemic species. By your definition Astatotilapia and many of the mbuna and malawi Haplochromines would be considered the same species.


Literature Cited:

-Barleunga, M., N. Stotting, W. Salzburger M Mishnick and A. Meyer (2006) Sympatric speciation in Nacaraguan crater lake cichlids. Nature, 7077, 19-23.

-Day, J.J., Santini, S., Garcia-Moreno, J., (2007) Phylogenetic relationships of the Lake Tanganyika cichlid tribe Lamprologini: the story from mitochondrial DNA. Mol. Phylogenet. Evol. 46, 629-642.

-Egger, B., Koblmüller, S., Sturmbauer, C., Sefc, K.M., (2007) Nuclear and mitochondrial data reveal different evolutionary processes in the Lake Tanganyika cichlid genus Tropheus. BMC Evol. Biol. 7, 137.

- Geiger MF, McCrary JK, Schliewen UK (2010) Not a simple case - A first comprehensive phylogenetic hypothesis for the Midas cichlid complex in Nicaragua (Teleostei: Cichlidae Amphilophus). Molecular Phylogenetics and Evolution, 56,1011-1024.

-Genner, M.J. & G.F. Turner (2012) Ancient hybridisation and phenotypic novelty within Lake Malawi’s cichlid fish radiation. Molecular Biology and Evolution 29, 195–206.

- Kobmuller, s., Egger, B., Sturmbauer, C., Sefc, K.M. (2010) Rapid radiation, ancient incomplete lineage sorting and ancient hybridization in the endemic Lake Tanganyika cichlid tribe Tropheini. Molecular Phylogenetics and Evolution, 55, 318–334.

-McKaye, K.R., Stauffer Jr., J.R., van den Berghe, E.P., Vivas, R., Lopez Perez, L.J., McCrary, J.K., Waid, R., Konings, A., Lee, W.-J., Kocher, T.D., (2002) Behavioral, morphological and genetic evidence of divergence of the Midas cichlid species complex in two Nicaraguan crater lakes. Cuadernos de Investigación de la UCA 12, 19–47.

-Sturmbauer, C., Salzburger, W., Duftner, N., Schelly, R., Koblmüller, S.(2010) Evolutionary history of the Lake Tanganyika cichlid tribe Lamprologini (Teleostei: Perciformes) derived from mitochondrial and nuclear DNA data, Molecular Phylogenetics and Evolution, 57, 266-284.
Befire we get swept into a morass of current phylogenetic studies and history of all Amphilophus , I think we should reflect on the other disciplines systemtatics values. Socrates and Thomas Aquinas believed there was a perfect heaenly model upon which all imperfect earth organisms were based. we have obfuscated the importance of such notions in the accurate descriptions of many of the earth's species by careful application of studies of morhology, behaviorin situ (I will not deign to answer another comment about fish interbreeding at home), ecology, and more recently genetics I applied these principles in my separation of two snakes from the genus Elaphe into Bogertophis, largely because they had a unque karyotype. I have no qualms with obvious genetic conclusions. This was cut and dry, 2 more chromosomes made you a new genus. I have applied the same principles to the Red Devil and -Midas cichlids where after careful review of all data available to me on genetic studies, Marta Barluenga's contention of indistinguish ability basically agreed with Seth Meek's contention that they appear to be the same species.

Very recently, there has been a contention that the proto-Amphilophus ancestors in L. Nicaragua had a propensity to develop thick lips. So apparently did human ancestors. This small difference in trait qualifies some humans as different races, and the same standards must be held for cichlids. In the body of my work, i elucidated all the minor differences that constite race in humans, and must therefore do so for cichlids.I recognize that ssytematics is subkjective, and some one is entitled to his opinion that it is a big enough difference to be a species. I agree they may some day be be isolated even within the Nicarguan Species Flock, as all evolution proceeds towards entropy. However, evolution has no set speed and the evidence is that Midas and Red Devils have been essentially the same during a time periods in which 15 to maybe 30 species nearby differentiated. Their interbreeding reinforces their similar appearance, and barring some major volcanic or diastrophic event which impacts the lake, I don't see change any time soon.
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Re: Proposed Sympatry of Midas and Red Devil Cichlids

Post by cichla » Thu Aug 09, 2012 2:27 am

Dear all.
RobertPrice, when I have read your post the first time I thought it must be a satire. But it seems to be that you are treating it serious. However, your comparison between the taxonomy and species discrimination of Midas Cichlids and Homo sapiens (in particular the shape and size of the lips) is just ridiculous.
In contrast to your statements about the taxonomy of the genus Homo there are many described species taxa. It starts already with Linneaus (1758) who recognized different species in genus. BasPels (see post above) explained why the taxonomy of the genus Homo has to be treated different.
Your thoughts about mate preferences and mate choice in Homo sapiens do not includes the knowledge and the results of the 'behavioral ecology' (which paint a much different picture). The species concept you are following is antiquated and just 'out of date'. Nowadays the majority of taxonomists (in particular ichthyologists) are convinced that species are evolving lineages. The 'modern' evolutionary species concept goes back to Simpson 1961 (see Wiley 1978) and that is much long before the rise of the 'DNA-stuff'.
You are classifying people ('greatest ichthyologist'; 'You seem expert on the fish themselves, but lacking...') instead of considering their papers, statements and arguments.
I agree with NuchalMan and Willem. Their contributions are well written and are including convincing arguments.
Best, IS
Wiley (1978): The Evolutionary Species Concept Reconsidered. Syst. Zool. 27: 17-26.
Last edited by cichla on Thu Aug 09, 2012 2:36 am, edited 1 time in total.

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