Reptile & Amphibian Forums

Moriarty and Cannatella (2004) write:

"The phylogenetic analyses of Cocroft (1994) and Da Silva (1997) are consistent with Hedges (1986) transferral of crucifer, regilla, cadaverina, and ocularis to a monophyletic Pseudacris. Our tree provides strong support for the monophyly of the genus including these taxa. Given that most checklists and field guides (Stebbins, 1985, is an exception) follow Hedges' taxonomy,
we continue it here."

Moriarty and Cannatella give no compelling reason for classifying crucifer, regilla, cadaverina and ocularis in Pseudacris. Morphologically most of these species are more like Hyla than Pseudacris in having well developed toe pads. Hyla cadaverina is an excellent climber. Even though Hyla regilla is more likely to be found on the ground than in trees in the wild, it too is an excellent climber. The genus Pseudacris, on the other hand, is an assemblage of secondarily terrestrial species with reduced toe pads. Immunological data also suggests that Hyla regilla, Hyla cadaverina and Hyla crucifer are basal to Pseudacris. Hyla crucifer is the species closest to Pseudacris phylogenetically. Hyla regilla and H. cadaverina both lie outside of the Pseudacris-H. crucifer clade. Therefore it is certainly possible to leave Hyla regilla and Hyla cadaverina outside of Pseudacris even if one were to insist on a holophyletic Pseudacris. Moriarty and Cannatella's decision to "follow most checklists and field guides" is a weak reason for maintaining a morphologically heterogeneous Pseudacris.

Stebbins' (1985) placement of regilla and cadaverina in Hyla is therefore justified on morphological grounds. "Most checklists" and field guides often blindly follow the latest taxonomic proposals. It appears that Stebbins has evaluated Hedges' taxonomic proposal and rejected it. I do not always agree with Stebbins' taxonomic decisions, but it appears that he does evaluate taxonomic proposals and in this case I think he has made the correct one in retaining these species in Hyla in the latest edition of his field guide. Further, immunological data by Maxson shows that Hyla cinerea, Hyla arborea and Hyla regilla form an unresolved polytomy. That means if Hyla regilla is included in Pseudacris, both Hyla arborea and Hyla cinerea may need to be included in Pseudacris as well, if one is using Moriarty and Cannatella's own Hennigian classificatory philosophy, which states that a taxon must include one ancestor and all of its descendants. However, if Hyla arborea is included in Pseudacris, it makes Pseudacris an invalid name, since Hyla arborea is the type species of Hyla and since Hyla is the earlier name, it has priority over Pseudacris under the rules of the ICZN. Moriarty and Cannatella have apparently ignored Maxson's immunological data but since their analysis does not include H. arborea, they cannot exclude the possibility that Hyla arborea and Hyla cinerea form a clade with Pseudacris, H. regilla, H. cadaverina and H. cinerea.

In sum, by including H. regilla and H. cadaverina in Pseudacris, Moriarty and Cannatella have not only created a morphologically heterogeneous group, with most members having reduced toe pads and a few having well developed toe pads, but they may have also created a potential can of taxonomic worms.

Reference

Moriarty, Emily C. and David. C. Cannatella 2004. Phylogenetic relationships of the North American chorus frogs (Pseudacris: Hylidae). Molecular Phylogenetics & Evolution. 30(2):409-420

You've got to be kidding. You're taking 26 year-old immunological data over a very well supported topology based on 2.4 kb of DNA? Not wise. Taxomic designations should be nontrivial, and you're taxonomic ideas clearly are. Only when taxonomic ranks are based on biological reality (evolutionary history, which is nontrivial) do they have maximum meaning and heuristic power.

There are some potential problems too with Moriarty and Cannatella's mtDNA data. They may have chosen a member of the ingroup as an outgroup. All of the species of Hyla found in the United States, according to immunological data, form a clade along with Acris, Pseudacris, L. ocularis and the Eurasian Hyla arborea species group. Taking both biogeography and morphology into consideration, it would appear that Hyla eximia is the ancestral species from which all other US species of Hyla evolved, given the fact that the bulk of its range is south of the border, the fact that the genus Hyla and the Hyla eximia species group is of neotropical origin, and the fact that practically all of the US species of Hyla plus Hyla arborea can be derived morphologically from an eximia-like ancestor. The "outgroup" chosen by Moriarty and Cannatella, namely Hyla chrysoscelis, Hyla andersonii and Hyla eximia, are all part of the same clade to which Pseudacris, H. regilla, H. crucifer, H. cadaverina and L. ocularis all belong. Their tree shows Hyla chrysoscelis, which is a derived member of this clade, to be the most basal! There is therefore the strong possibility that some of the apomorphs found in H. chrysoscelis have been mistaken as plesiomorphs for this clade.

Hence their analysis is far from the final word on the relationships among members within this group, especially since immunological data shows that there was a veritable adaptive radiation very early in the history of this group's existence, as Maxson's 1976 paper shows. Moriarty and Cannatella could have, and should have, chosen as outgroup another species that is more distantly related to the North American species of Hyla than H. chrysoscelis and Hyla andersonii. Smilisca baudini or Osteopilus septentrionalis, for example, may have been better choices as outgroup than Hyla andersonii and Hyla chrysoscelis, although I would have left Hyla eximia in the analysis, given the possibility that it is the ancestral species of all US Hyla and their closest relatives.

but you haven't shown that any of their conclusions are wrong. Their finding were in concordance with Hedges (1986) and da Silva's (1997) findings. You suggest that "Moriarty and Cannatella should have chosen as outgroup another species that is more distantly related to the North American species of Hyla than H. chrysoscelis and Hyla andersonii." That's baloney. Choosing very distant taxa as outgroups does not polarize characters nearly as well as as closely related taxa. While their discussion of outgroup selection was lacking, they did base it on prior findings. The relationships they found within Pseudacris are fine, the question remains on the position of this clade within Hylidae. You claim Hyla chrysoscelis is most more derived, but relative to what? What's your source here? And again, the resolving power of that immunological data cannot compete with their mtDNA data.

As I said, it is most likely that Hyla eximia is the most basal member of US Hyla and their closest relatives, given its geographic location and morphology and given the virtual certainty that the genus Hyla originated in the neotropics (vs. for example the nearctic or palearctic). Moriarty and Cannatella's tree surprisingly shows Hyla chrysoscelis as the most basal member, which is problematic, since 2 of Hedges' 3 trees show that Hyla eximia is the most basal species (fig. 1a and 1b of Moriarty and Cannatella). Further, Hyla crucifer has traditionally been considered a species of Hyla on the basis of morphology. Yet their maximum likelihood tree has H. crucifer nested deeply within Pseudacris, a group of degenerate hylids with reduced toe pads. That placement is not parsimonious, since it requires an evolutionary reversal for H. crucifer to have descended from an ancestor with reduced toe pads. Their maximum parsimony tree shows that H. crucifer forming an unresolved polytomy with 2 other groups of species traditionally placed in Pseudacris. Moriarty and Cannatella's parsimony tree therefore seems to make more sense than their likelihood tree. Their parsimony tree is also closer to the immunological data, which you dismiss casually because of its age.

According to immunological data, Hyla crucifer is basal to Pseudacris and Hyla regilla and Hyla cadaverina are in turn basal to H. crucifer. da Silva's tree also shows the same topology and both of these contradict Moriarty and Cannatella's likelihood tree. Using da Silva's tree, one can certainly recognize a holophyletic Pseudacris while leaving Hyla cadaverina, H. regilla and H. crucifer outside of Pseudacris. Hyla is paraphyletic whether we accept Moriarty and Cannatella's classification or not. Therefore, why create heterogeneity within Pseudacris by including several treefrogs that have been traditionally classified within Hyla? Why do so if there is a danger of having Pseudacris invalidated? Moriarty and Cannatella's reason for doing so (i.e. they are following most checklists and field guides) is dubious. The reason why Pseudacris has been recognized as being distinct from Hyla is the reduced toe pads of this group of secondarily terrestrial hylids. By including Hyla crucifer, Hyla regilla and H. cadaverina, all of which have well developed toe pads, the genus Pseudacris becomes undefinable and undiagnosable, notwithstanding Hedges' claim that his "Pseudacris" has a "cold-weather breeding season, a round or ovoid testis, and a black pigment covering on the testis." Breeding season is not a taxonomic character. The shape of the testis and the presence of dark pigment only apply to the males of these species. The reduced toe pads are diagnostic of both sexes.

To start, you and I have different philosophies regarding taxonomy and monophyly. While I can recognize paraphyletic species, I believe any designation genus or above should be monophyletic. I know you believe differently, and I don't want to waste time arguing about something like that when neither of us will change our minds.
And regarding me "casually dismissing" immunological data, that is a total misrepresentation. I stated that the resolving power of dna sequence data is superior to that of immunological data. Additionally, that remark was made referring to your unresolved tritomy.
If P.crucifer were to be recovered basal to the other two clades in the unresolved parsimony tree (not super likely, but theoretically possible), you would have a point about not needing to include P.crucifer, P. cadaverina or P.regilla in Pseudacris since Hyla doing so would not render Hyla monophyletic either. Ultimately there needs to be a large analysis, which Cannatella has a grant to do.
Regarding the placement of P. crucifer not being parsimonious because it would necessitate a reversal (or multiple losses of toe pads), that's should not be a deciding factor. Nature would have no problem preforming either.

Yes Moriarity, Cannatella and you differ from I in taxonomic practice. Specifically the three of you are following Hennig and are intolearant of paraphyletic taxa. But as I pointed out, whether or not one includes Hyla regilla, Hyla crucifer and Hyla cadaverina in Pseudacris, Hyla will be paraphyletic, as Duellman pointed out. Pseudacris is not paraphyletic even if these species are excluded. Although Hedges thought that these species must be included to maintain a holophyletic Pseudacris, and it was indeed the reason for his original taxonomic proposal, both da Silva and Moriarty and Cannatella's data suggest that Hyla regilla and Hyla cadaverina can be excluded without affecting the holophyly of Pseudacris. Therefore it boils down to whether one should include these species in Pseudacris because "most checklists and field guides" included these species, as Moriarty and Cannatella stated. As I have already stated, Hyla arborea is closely related to Hyla regilla Hyla eximia and Hyla cinerea, so close that if Hyla regilla is included in Pseudacris, Hyla arborea may need to be included to maintain the holophyly of Pseudacris. But if Hyla arborea is included, then Pseudacris is a junior synonym of Hyla since Hyla has priority over Pseudacris, and since Hyla arborea is the type species of Hyla. Further, why does one recognize Pseudacris in the first place? Pseudacris has been recognized traditionally because it is a monophyletic group and because it is a group of degenerate, secondarily terrestrial hylids with reduced toe pads. The inclusion of Hyla regilla and Hyla cadaverina in Pseudacris destroys this definition and makes Pseudacris undiagnosable. Hedges, in redefining Pseudacris, found that his "Pseudacris" are frogs with a "cold-weather breeding season, a round or ovoid testis, and a black pigment covering on the testis." As I pointed out, breeding season is not a taxonomic character and the morphology of the testis apply only to the males of these species. It turns out that Hedges' redefinition is unnecessary according to da Silva's data; Pseudacris can indeed be a holophyletic group even if Hyla regilla and Hyla cadaverina are excluded. So, why insist on including these species, if their inclusion creates a heterogeneous Pseudacris and at the same time opens the possibility that Pseudacris will become a junior synonym of Hyla? Moriarty and Cannatella appears to be attempting to maintain some semblance of taxonomic stability in their decision to leave Hyla regilla and Hyla cadaverina in Pseudacris. By doing so, however, Moriarty and Cannatella may have opened the door to the inclusion of Hyla arborea in Pseudacris; they may have unwittingly opened a can of taxonomic worms.

Your claim that immunological data has less resolving power than newer techniques is certainly true. mtDNA data does have a lot more resolving power. However, Moriarty and Cannatella's parsimony tree still shows an unresolved polytomy, suggesting that there may have been an adaptive radiation. Immunological data suggests the same in the early history of this group of holarctic Hyla. Besides, despite the higher resolving power of mtDNA, mtDNA can still be less informative if one is not careful about the choice of outgroup. Moriarty and Cannatella's choice of outgroup species (Hyla chrysoscelis) is analogous to choosing Pan as the "outgroup" when one is analyzing the relationships among Gorilla, Homo, and Pongo. They are in fact using a derived member of the ingroup as the "outgroup" species.

Moriarty and Cannatella's likelihood tree, as I said, requires a reversal. Either Hyla crucifer re-evolved large toe pads from an ancestor with reduced toe pads, which is unparsimonious, or as you suggest "multiple loss of toe pads." That means the species traditionally classified under Pseudacris independently lost their toe pads. This scenario is even less parsimonious since Pseudacris is a monophyletic group. It is true that multiple independent loss or reacquisition of a lost character are both possible, but they are less probable than the alternative hypothesis, especially since immunological data and da Silva's data both support the basal position of Hyla crucifer relative to Pseudacris, making the ad hoc hypothesis of multiple independent loss or reacquistion unnecessary.

In sum, there are at least four reasons for excluding Hyla regilla, Hyla cadaverina and Hyla crucifer from Pseudacris:

1. problems with Moriarty and Cannatella's analysis because of their unfortunate choice of outgroup species,

2. the availability of more than one set of data suggesting that Hyla regilla, Hyla cadaverina and Hyla crucifer can be excluded from Pseudacris without affecting its holophyly

3. inclusion of these species creates a heterogeneous Pseudacris

4. inclusion of these species may endanger the validity of Pseudacris because Hyla arborea (the type species of Hyla) is closely related to Hyla eximia, Hyla regilla, Hyla chrysoscelis et al.

1. The immunological data tree from '95 doesn't indicate there's a problem with outgroup selection.
2. The best available data do not support P.crucifer falling outside other Pseudacric species.
3. What is a heterogeneous Pseudacris? Some having toe pads and other not? Not a big deal. They share a common evolutionary history and that's what's important.
4. The '95 immunological data give no indication of the possibility of H.arborea falling within Pseudacris.

1. Sorry for the typo in the original post. The tree was from the 1978 paper, not 1995. Maxson and Wilson originally published their data in 1975, in Syst. Zool. In that paper, they found H. chrysoscelis, Hyla eximia, and Hyla femoralis forming an unresolved polytomy with the lineage leading to Hyla gratiosa and Hyla cinerea. This clade in turn forms an unresolved polytomy with the branch which leads to Hyla crucifer, Pseudacris triseriata and Hyla regilla. Hence there is indeed data which suggests that Moriarty and Cannatella's choice of outgroup species (Hyla eximia, Hyla andersonii and Hyla chrysoscelis) is problematic, since Hyla chrysoscelis and Hyla eximia are part of the ingroup, not an outgroup.

2. The best available data does suggest that Hyla crucifer is outside of Pseudacris. In terms of morphology, it is not a Pseudacris, according to most systematists who rely on morphological data, including William Duellman himself. Further, da Silva's data does show H. crucifer outside of traditional Pseudacris.

3. If a "common evolutionary history" is all that matters, there is no compelling reason for including Hyla regilla and Hyla cadaverina in Pseudacris since Hyla andersonii, Hyla eximia, and Hyla chrysoscelis also share a common evolutionary history with Pseudacris and thus these too can be included. If heterogeneity is not a concern, then why bother recognizing Pseudacris in the first place, why not synonymize this group of degenerate hylids with Hyla? As Duellman pointed out in his revised edition of the Hylid Frogs of Middle America, Hyla is paraphyletic if Pseudacris is recognized. Apparently Moriarty and Cannatella thought that Pseudacris is deserving of taxonomic recognition as a genus, even if it renders Hyla paraphyletic. Why is Pseudacris deserving of such recognition? Is it because it is a clade? Well, if so, then one can certainly include Hyla eximia and H. aborea in this clade as well. Of course including Hyla arborea in this clade will turn Pseudacris into a junior synonym of Hyla, given the fact that Hyla arborea is the type species of Hyla and the fact that Hyla has priority over Pseudacris.

4. If you would take the trouble to read Maxson and Wilson's 1975 paper, you should be able to see that the branch that leads to Hyla regilla, Hyla crucifer and Pseudacris triseriata forms a polytomy with 3 other branches, one of which includes Hyla arborea. Therefore the same ancestor that gave rise to Pseudacris and Hyla regilla is also ancestral to Hyla arborea. Under the rules of Hennigian taxonomy, the ancestor of Pseudacris, Hyla crucifer and Hyla regilla and all of the descendants of this ancestor must be included in the same taxon (in this case genus Pseudacris). If so, then Hyla arborea must also be included, since it shares the same ancestor as Pseudacris, Hyla crucifer and Hyla regilla. That would mean that Hyla and Pseudacris is the same genus, which in turn means, once again, that Pseudacris is a junior synonym of Hyla.

As I pointed out before, Moriarty and Cannatella did not include Hyla arborea in their analysis. Therefore they cannot refute Maxson and Wilson's data. Unless and until Maxson and Wilson's immunological data has been refuted, Moriarty and Cannatella's "Pseudacris" is indeed a junior synonym of Hyla according to Hennigian taxonomy (which is being practiced by Moriarty and Cannatella) and the rules of the ICZN.

1. I have the Maxson and Wilson (1975) paper in hand, and there is no indication whatsoever that Moriarty and Canatella's choice of outgroups is problematic. All three species clearly fall outside the monophyletic clade they are investigating. I would suggest your understanding of systematics is flawed.
2. The best data are Moriarty and Canatella's MP and ML trees. Maximum likelihood generally outperforms parsimony, so I would say the ML tree is to be most trusted. But neither tree supports P. crucifer being basal to other Pseudacris. However, it could be tested statistically whether the placement of crucifer as basal to others results in a statistically worse tree. While morphology can be useful in deciferring relationships, your use of one character (toe pads) that you arbitrarily choose is hardly scientific.
3. Yes, Hyla is paraphyletic if Pseudacris and Acris are recognized. Are you suggesting those genera should be sunk? Again, your discussion of H.arborea is irrelevant; it has nothing to do with Pseudacris. (No, it is not an ingroup!) Pseudacris (including regilla and crucifer) share a common ancestor that H. arborea clearly does not.
4. Again, you're wrong with H. abrorea (per Maxson and Wilson 1975). It is clearly an outgroup of the taxa Moriarty and Canatella are addressing.
Moriarty and Canatella aren't refuting Maxson and Wilson's findings; in fact, they're congruent! Moriarty and Canatella are addressing relationships within a clade that Maxson and Wilson found monophyletic, even in the conservative tree (you failed to mention the resolved tree). The recent paper has increased taxon sampling and uses DNA sequence data. If the new study were paired down to only the taxa that Maxson and Wilson included, it would result in the exact same topology (so far as Psuedacris is concerned, which, not surprisingly, the recent paper is concerned about). Dude, I just don't understand where you're coming from.

1. You are mistaken. The "Pseudacris" of Moriarty and Cannatella clearly shares a common ancestor with Hyla arborea, Hyla eximia and Hyla chrysoscelis according to Maxson (1978, J. Herpetology). Hyla eximia and Hyla chrysoscelis are clearly part of the ingroup, not an outgroup according to Maxson and Wilson (1975). There is nothing wrong with my understanding of systematics. You are simply dogmatic.

2. On what basis do you claim that Moriarty and Cannatella's data are the best? Their ML and MP trees contradict each other. Since there is only one version of history, one or both of their tress is definitely wrong. The ML tree shows Hyla crucifer evolving from an ancestor with reduced toe pads, which is clearly unparsimonious. The alternative, as I pointed out before, is the multiple independent loss of toe pads among the members of Pseudacris, which would clearly be even less parsimonious. It also means that this character is a homoplasy and therefore this clade would be undeserving of recognition since it would be a polyphyletic group. Both ML and MP trees also show the surprising placement of Hyla chrysoscelis being the most basal species, which is contradicted by Maxson and Wilson's immunological data.

3. If Hyla is paraphyletic no matter how we draw the line that separates Pseudacris from Hyla, then why do we draw a line in such a way that includes Hyla crucifer, Hyla cadaverina and Hyla regilla in Pseudacris, when doing so destroys the classic definition of Pseudacris? Why not exclude Hyla crucifer, Hyla regilla and Hyla cadaverina, since Pseudacris would still be holophyletic without these species? If one relies on Moriarty and Cannatella's MP tree, and if one exchanges the positions of the Hyla crucifer branch and the "Fat Frogs" branch, then one can easily see that Hyla crucifer, Hyla regialla and Hyla cadaverina can be left out of Pseudacris without upsetting Hennigians like you. Indeed, by claiming that they are only following "most checklists and field guides" in leaving these species in Pseudacris, Moriarty and Cannatella are tacitly admitting that they could have taken these species out without violating Hennigian dogma.

4. If you are correct that Moriarty and Cannatella's data are congruent with Maxson and Wilson's paper, then why can't they classify Hyla regilla, Hyla crucifer and Hyla cadaverina in Hyla, since Maxson and Wilson did just that? Why should any taxonomist prefer a classification that results in a heterogenetous Pseudacris? Why recognize Pseudacris at all if the inclusion of several hylids with well developed toe pads renders it difficult to define? I object to Moriarty and Cannatella's classification because it results in a heterogeneous Pseudacris that is at the same time difficult, if not impossible, to define. Lastly, Moriarty and cannatella's trees, which show Hyla chrysoscelis as the most basal species, is clearly contradicted by Maxson and Wilson's data, which shows H. chrysoscelis to be a highly derived species of the Holarctic hylid clade. In fact, Maxson and Wilson's data clearly is better, since they picked the correct outgroup, unlike Moriarty and Cannatella.

First, Linda Maxson is the dean of my college and will be the first to tell you that the techniques used in her earlier papers have fallen by the wayside for better, more modern techniques. Just follow her paper trail.
Second, those who can, do, those who can't, criticize.

Maxson would be the last person on earth to tell anyone, including you, that her data has "fallen by the wayside for better, more modern techniques." It is true that immunological techniques are no longer used widely today, because of the ease with which DNA can be sequenced, but the molecule that Maxson and Wilson were working with, namely serum albumin, is highly informative, because it is not constrained by natural selection. Amino acid substitutions within this molecule are therefore due purely to chance, and it is therefore an excellent molecule for estimating relative lineage divergence times. Systematists have rarely found a better molecular characters with which to work. Even mtDNA mutations are not completely neutral as it is still very much functional, unlike serum albumin.

Secondly, only those who are dogmatic, for example the cladists, would openly ignore criticism and/or opposing evidence. There is no shortage of evidence opposing the placement of Hyla chrysoscelis as the most basal species in Moriarty and Cannatella's tree, and there is no shortage of criticism on the cladists' "destructive, impractical and scientifically untenable" classifications (Mayr and Ashlock 1991).

from her CV:

Hay, J. M., I. Ruvinsky, S. B. Hedges, and L. R. Maxson. 1995. Phylogenetic relationships of amphibian families inferred from DNA sequences of mitochondrial 12S and 16S ribosomal RNA genes. Molec. Biol. Evol. 12: 928-937.
Lopez, T. J., and L. R. Maxson. 1995. Mitochondrial DNA sequence variation and genetic differentiation among colubrine snakes (Reptilia: Colubridae: Colubrinae). Biochem. Syst. Ecol. 23: 487-505.
Hedges, S. B., and L. R. Maxson. 1996. Molecules and Morphology in Amniote Phylogeny. Molec. Phylogenetics Evol., 6:312-314.
Ruvinsky, I., and L. R. Maxson. 1996. Phylogenetic relationships among Bufonoid frogs (Anura: Neobatrachia) inferred from mitochondrial DNA sequences. Molec. Phylogenetics Evol. 5:533-547.
Lopez, T. J., and L. R. Maxson. 1996. Albumin and mitochondrial DNA evolution: Phylogenetic implications for colubrine snakes (Colubridae: Colubrinae). Amphibia/Reptilia 17: 247-259.
Roberts, J. D., P. Horwitz, G. Wardell-Johnson, L. R. Maxson, and M. J. Mahony. 1997. Taxonomy, relationships, and conservation of a new genus and species of Myobatrachid frog from the high rainfall region of Southwestern Australia. Copeia, 1997(2): 373-381.
Futuyma, D.J., R.R. Meagher, M.J. Donoghue, J. Hanken, C.H. Langley, and L. Maxson. 1999. Evolution, Science, and Society: Evolutionary Biology and the National Research Agenda: http://www.amnat.org
Hass, C. A., M. DiRado, L. R. Maxson, and S. B. Hedges. 1999. Relationships and biogeography of West Indian hylid frogs: an analysis of serum albumin transcript sequences. Joint meeting of ASIH, HL, and SSAR. Penn State. 6/29/99.
Hass, C. A., L. R. Maxson, and S. B. Hedges. 2001. Relationships and divergence times of West Indian amphibians and reptiles: Insights from albumin immunology. Pp. 157-174. In C. A. Woods & F. E. Sergile (eds), Biogeography of the West Indies: Patterns and Perspectives. 2nd edition, CRC Press.
Maxson, L.R. 2001. Micro-Complement Fixation (MC’F). In Encyclopedia of Genetics. W. A. Fitch et al., eds. Academic Press.

Note also that most of the sero-albumin work she has done of late is in CONJUNCTION with mtDNA work . . . the thing is, mtDNA and microsat nuclear DNA simply give so many more characters to evaluate than sero-albumin . . .

Troy

"Hass, C. A., M. DiRado, L. R. Maxson, and S. B. Hedges. 1999. Relationships and biogeography of West Indian hylid frogs: an analysis of serum albumin transcript sequences. Joint meeting of
ASIH, HL, and SSAR. Penn State. 6/29/99.
Hass, C. A., L. R. Maxson, and S. B. Hedges. 2001. Relationships and divergence times of West Indian amphibians and reptiles: Insights from albumin immunology. Pp. 157-174. In C. A. Woods
& F. E. Sergile (eds), Biogeography of the West Indies: Patterns and Perspectives. 2nd edition, CRC Press.
Maxson, L.R. 2001. Micro-Complement Fixation (MC’F). In Encyclopedia of Genetics. W. A. Fitch et al., eds. Academic Press.

Note also that most of the sero-albumin work she has done of late is in CONJUNCTION with mtDNA work . . . the thing is, mtDNA and microsat nuclear DNA simply give so many more characters to evaluate than sero-albumin . . ."

Troy cited a large number of papers by Linda Maxson, in which mtDNA was used. However, he has also noticed that her most recent writings dealt with serum albumin. That supports my earlier assertion that immunological data has not been supplanted by more recently developed techniques and my assertion that Maxson would be the last person on earth to suggest that her immunological data should be discarded or ignored. As I pointed out, the ease with which DNA can be sequenced makes immunological techniques unattractive to the new generation of systematists, especially since the older technique often requires the sacrifice of a live animal if it is small, whereas a small tissue sample from a roadkill or even a preserved specimen is all that is needed with some of the newer techniques. mtDNA has even been extracted from the skull of a Neanderthal.

But as I pointed out earlier as well, serum albumin is a neutral molecule in which amino acid substitutions are unconstrained by natural selection. As such it is an excellent molecule with which to estimate relative lineage ages and timing of divergence events, since amino acid substitutions and the underlying nucleotide substitutions are more clock like than most molecules, even mtDNA, since mtDNA is still a functional molecule. As one can see from the last few papers listed above, immunological data was used for estimating lineage age and divergence times.

Unfortunately, Troy has very little understanding of the advantages of using neutral characters to determine phylogenetic relationships. Hence he thinks he can simply ignore Maxson's immunological data. It is not a good idea to ignore scientific data. Troy is apparently trying to justify dismissing immunological data by showing us that Maxson herself has turned to mtDNA in some of her recent studies. What Troy fails to understand is that mtDNA data does not invalidate immunological data any more than the invention of an electrical nailing gun makes a house built with a hammer a piece of junk. The electrical nailer simply makes buiding houses faster; its use does not guarantee the building of a better house. Similarly, mtDNA data is not necessarily better than immunological data. In some cases, it can even be worse, if one is not careful about choosing an outgroup for example. Immunological techniques and mtDNA sequencing are both tools for the systematist. In the hands of a knowledgeable systematist, either of these tools can be equally effective in constructing a sound phylogeny. In the hands of an inexperienced systematist, they can produce anomalous results.

Of course, it also takes knowledge to be able to judge the results of a systematic analysis. Those who lack the ability to judge the results and/or conclusions of a systematic analysis will simply turn to the Troy method: simply accept the newest data as the most accurate because they must be the best if they are using the latest techniques. Life would be too simple (and boring) for the systematist if the Troy method works, since all one has to do is to obtain a nose ring and attach it to the Troy method. Unfortunately, the Troy method does not work. Life is therefore not simple for the systematist. Nor is life boring for the knowledgeable systematist.

he now resorts to personal attacks every time he responds to me . . .

by the way, how is an area which receives greater than 10" or rainfall per year still considered a desert? Deserts are, by definition, areas that receive 10" of rainfall or less per year . . .

LOL

Troy

I am sorry to see that Troy has failed to respond to any of my points in the lengthy rebuttal to his claim that Linda Maxson has now been using mtDNA data, and which ostensibly means that immunological data should be ignored.

Sorry, Troy, immunological data is scientific data. It is not a good idea to ignore scientific data, unless you are a dogmatic person. Are you a dogmatic person?

As for Troy's claim that the Chihuahuan Desert is not a desert, he has backed down from that ridiculous claim and now calls West Texas part of the Chihuahuan Desert "biotic province." A desert biotic province is a region where the biota (plants and animals) is adapted to desert conditions. That means the plants and animals living in West Texas are desert plants and desert animals. Desert plants and desert animals live in deserts. Apparently Troy has proven these desert plants and desert animals "wrong" because they have no rights to live in a place where they are not supposed to, since Trans Pecos Texas is not a desert. The next logical step perhaps is for these plants and animals to admit that they are wrong, and to move out of West Texas. Troy has a lot of persuasion in front of him, because there does not seem to be a mass exodus of desert plants and desert animals from West Texas. They are not listening to Troy. They continue to live in West Texas as though it were a desert.

No, you do not understand systematics. Your comparisons of Maxson and Wilson's study with that of Moriarty and Cannatella's are flawed. They are not addressing the same question; the later study is only looking at a single clade within Maxson and Wilson's. For what Moriarty and Cannatella attempt to do (address relationships within Pseudacris), there outgroups are fine. They're not attempting to resolve relationships among all Hyla. And the polytomy in the MP tree does not contractict the topology of the ML tree.

1. "[Moriarty and Cannatella] are not addressing the same question [as Maxson and Wilson]

me:
That is partly true. Maxson and Wilson were primarily looking at relationships among the holarctic Hyla, although they did include Acris, Pseudacris and Limnaodis in their analysis. Moriarty and Cannatella are primarily looking at whether Pseudacris is a holophyletic group. Both Maxson and Wilson's immunological data and Moriarty and Cannatella's analysis nevertheless show that Hyla crucifer is the closest relative of Pseudacris. Moriarty and Cannatella's ML tree requires the unlikely (though not by any means impossible) scenario of Hyla crucifer re-evolving well developed toe pads from an ancestor with reduced toe pads. Their MP tree is in agreement with Maxson and Wilson's data in that both show Hyla crucifer as the sister species of Pseudacris. Given the fact that da Silva also shows a similar topology, it is more likely than not that Moriarty and Cannatella's ML tree is incorrect.

2. "[Moriarty and Cannatella's] outgroups are fine."

me:
Once again I would have to disagree with that assertion. If one looks carefully at Maxson and Wilson's data, theirs show the Hyla regilla/H. crucifer/Pseudacris branch forming an unresolved polytomy with the Hyla eximia/Hyla chrysoscelis branch, and the Hyla arborea branch. That means the common ancestor of Hyla regilla, Hyla crucifer and Pseudacris is also the common ancestor of Hyla arborea, Hyla eximia and Hyla chrysoscelis. Moriarty and Cannatella's "Pseudacris" is defined cladistically as the common ancestor of Hyla regilla, Hyla crucifer and Pseudacris (traditionally defined) plus all of the desendants of this ancestral species. Using that definition, all of the species of Hyla found in the United States , in Europe (including Hyla arborea) and in Asia thus belong to the genus Psuedacris. That makes Pseudacris a junior synonym of Hyla.

The question remains: why would anyone want to include Hyla regilla, Hyla crucifer and Hyla cadaverina in Pseudacris if by doing so Pseudacris becomes undefinable and so heterogeneous as to be useless, and if doing so means that this genus is now no longer a valid name under the rules of the ICZN?

you:
If one looks carefully at Maxson and Wilson's data, theirs show the Hyla regilla/H. crucifer/Pseudacris branch forming an unresolved polytomy with the Hyla eximia/Hyla chrysoscelis branch, and the Hyla arborea branch. That means the common ancestor of Hyla regilla, Hyla crucifer and Pseudacris is also the common ancestor of Hyla arborea, Hyla eximia and Hyla chrysoscelis. Moriarty and Cannatella's "Pseudacris" is defined cladistically as the common ancestor of Hyla regilla, Hyla crucifer and Pseudacris (traditionally defined) plus all of the desendants of this ancestral species.
me:
No, the fact that Pseudacris (including crucifer and regilla) form their own exclusive branch in all analyses (including Maxson and Wilson's), something that you acknowledge above, means that they share a more recent common ancestor exclusive of the one uniting all Hyla.
Look at taxon sampling in Moriarty and Canatella relative to Maxson and Wilson. That is the reason you "H. crucifer" is sister to Pseudacris, not nested within.

"No, the fact that Pseudacris (including crucifer and regilla) form their own exclusive branch in all analyses (including Maxson and Wilson's), something that you acknowledge above, means that they share a more recent common ancestor exclusive of the one uniting all Hyla."

You would be correct if new species evolve at regular and short intervals. It is true that the node from which Hyla regilla, Hyla crucifer and Pseudacris branch off does not coincide exactly with the node from which Hyla arborea and Hyla eximia/chrysoscelis branched off. The immunological distance between those two nodes, however, is short. In 1974, Maxson and Wilson thought that Hyla eximia is only convergently similar to Hyla regilla morphologically because these two are separated by a surprising amount of immunological distance. In 1975 they probably rethought their hypothesis after they found that these two also shared a common ancestor with Hyla arborea, which is also similar morphologically to Hyla regilla and Hyla eximia. Hyla regilla, Hyla arborea and Hyla eximia are almost certainly morphologically conservative species that resemble each other because they have changed little and because they resemble their common ancestor. That common ancestor, which is also the ancestor of all holarctic hylids is most likely Hyla eximia, given its morphology and biogeography. Evolutionary stasis, which is probable in a morphologically conservative lineage, may have kept Hyla eximia unchanged even though numerous species have budded off of it. The species from which all holarctic hylids evolved is probably the same species as the one from which Hyla regilla, Hyla crucifer and Pseudacris evolved.

"Look at taxon sampling in Moriarty and Canatella relative to Maxson and Wilson. That is the reason you "H. crucifer" is sister to Pseudacris, not nested within."

I disagree. It is instead the choice of outgroup species which accounts for the anomalies one sees in Moriarty and Canatella's ML and MP trees. Both trees surprisingly show H. chrysoscelis more basal to Hyla eximia. As I pointed out above, Hyla eximia is most likely the ancestor of all holarctic hylids (judging from biogeography and morphology) and it is also morphologically conservative (as I explained above). If Hyla chrysoscelis is more basal, that means Hyl eximia probably evolved from a Hyla chrysoscelis morphotype. But since there are several species that are morphologically similar to Hyla eximia in Mexico (namely members of the Hyla eximia group as defined by Duellman), Moriarty and Cannatella's tree would require these species (which are morphologically close to Hyla eximia) to have evolved convergently to resemble Hyla eximia, or it would require Hyla chrysoscelis to have evolved from an ancestor that looked like Hyla eximia, and then Hyla eximia would evolve by undergoing a reversal from a H. chrysoscelis-like ancestor. In other words, in order to embrace Moriarty and Cannatella's tree topology, one has to invoke multiple unparsimonious scenarios, not only in the evolution of toe pads but also in the evolution of the Hyla eximia morphotype. Personally I would not invoke such a large number of ad hoc hypotheses in order to embrace Moriarty and Cannatella's phylogeny, nor would I embrace their classification since it is both heterogeneous and probably unstable due to the possibility that the inclusion of Hyla arborea in their "Pseudacris" is unavoidable.

For the record, DHL stopped responding to my posts when I pointed out that Hyla eximia is most likely the ancestral species and/or morphotype from which all holarctic hylids evolved, given its biogeography and the morphological similarity between Hyla regilla, Hyla arborea and Hyla eximia and their large immunological distances. I thought that DHL had disappeared from this forum but he proved that he did not when he responded instead to Patrick Alexander's off topic personal attack, which alleges that I somehow reject data from Hennigians and that I also reject data that are published since the 1980's. I have since shown that Patrick Alexander's claims are baseless and easily contradicted in a different part of this thread.

Returning to the problem of classifying Hyla regilla, it is also possible that Hyla regilla was the species that evolved out of the node from which Hyla arborea also evolved. This is supported by immunological data by Maxson and Wilson, published in 1974 in the journal Science, which shows that some island populations of Hyla regilla are almost as distant immunologically from mainland populations of Hyla regilla as they are from Hyla eximia. Such data shows that Hyla regilla is an old species, and that it is morphologically conservative. Cladistic ideology assumes that an ancestor automatically becomes extinct when new species arise. But evolutionary biologists who have studied speciation extensively know that new species often form by the process of budding, or peripatric speciation. That means a small isolated population evolved into a new species while the ancestral species continues to live largely unchanged. If Pseudacris and Hyla crucifer had evolved by budding off of Hyla regilla, that would mean that the entire Hyla regilla-Pseudacris-Hyla crucifer branch is in fact Hyla regilla. Since this branch shares the same ancestor as Hyla arborea and Hyla eximia, putting Hyla regilla in Pseudacris also mean that Hyla arborea and Hyla eximia must also be transferred into Pseudacris in order to avoid making Pseudacris a paraphyletic genus. DHL contends that Moriarty and Cannatella's mtDNA data is the best available. I asked him/her how he/she made that determination but received no response. Whether or not Moriarity and Cannatella's data is the best available, the plain fact is that Moriarty and Cannatella did not include Hyla arborea in their analysis. They therefore cannot refute Maxson and Wilson's immunological data. That means their analysis cannot be used to show that Pseudacris is a valid genus that excludes Hyla arborea. Further studies are needed to clarify this point, but the available data does indeed show that the inclusion of Hyla regilla (and Hyla cadaverina) in Pseudacris is an unwise idea taxonomically. Of course their inclusion also makes Pseudacris too heterogeneous to be meaningful, notwithstanding Moriarty and Cannatella's claim that this genus can be redefined on the morphology of their testes.

Since R. C. Stebbins' new field guide excludes both H. regilla and H. cadaverina from Pseudacris, there should be no rush to adopt Moriarty and Cannatella's taxonomic arrangement, since they themselves are merely following checklists and field guides. Stebbins' arrangement also makes more sense because the traditional definition is that Pseudacris is a monophyletic group characterized by reduced toe pads, which are the result of a secondarily terrestrial existence. Excluding Hyla regilla and Hyla cadaverina from Pseudacris preserves this definition without making Pseudacris any less of a monophyletic group. Only those who like to make classifications so heterogeneous as to be useless would stubbornly insist on including Hyla regilla and Hyla cadaverina in Pseudacris.

The problem is that you don't understand systematics. And I'm not trying to be confrontational or rude about it, but you don't. I stopped responding because this is going nowhere. We are so far from being on the same page that this is pointless. But there are clearly no data that would support the postition of H.eximia or H.arborea being nested within Pseudacris. By the way, it is very odd how you always resort to discounting someone by calling them Hennigian, even though the argument has nothing to do with that. I'm done with this thread.

You have made the assertion that I do not understand systematics, but you also asserted that Moriarty and Cannatella's data are the best available. I refuted your claim by pointing out that

1. there is only one version of history and the MP tree of Moriarty strongly conflicts with their ML tree. Therefore one or both of their trees are wrong.

2. it is unparsimonious for Hyla chrysoscelis to be basal to Hyla eximia, but both the ML and the MP trees of Moriarty and Cannatella show this anomaly. 2 of Hedges' 3 trees, on the other hand, show Hyla eximia basal to Hyla chrysoscelis and so does da Silva's tree, which was reproduced in Duellman (2001).

There is indeed immunological data to support that Hyla arborea is nested within "Pseudacris," if Hyla regilla and Hyla crucifer are included in Pseudacris. The "Pseudacris" branch shares a common ancestor with Hyla arborea and Hyla cinerea according to the tree below, which is reproduced with some modifications from Maxson (1978). Just replace Hyla regilla with Pseudacris in that tree and you will see why Pseudacris will instantly become a junior synonym of Hyla. Notice that I did not mention anything about Hennig or Henngian above to dismiss your ideas. I used nohting but data to falsify your claims.

That said, Hennigians do have the unfortunate habit of ignoring facts, especially those that clash with their ideology. And I have proven that here as well.

And this one is harder for you to do the word-twister with! LOL

wild-caught specimen from Jeff Davis County, TX . . .

In response to Troy, I will only simply quote what I said in the other forum, since Troy had no answer to what I said there in response to his picture of a snake without the silvery gray iris.

Me:
"I will not, for the time being, contest troy h's claim that someone has collected an alterna without a silvery gray iris. I will instead try to figure out what such a find, if it were authentic, would do to my theory of how L. mexicana transformed into L. alterna.

Of course, if it were a fake, then it would not change my theory. Assuming, however, that it is real, I will see what effects it will have on my theory.

1. It is a mutant. Mutations happen sooner or later. If it is a mutant then it has no effect on my theory. In fact, it supports my contention that the silvery gray iris (or iris color in general) in L. alterna is a selectively neutral character that became fixed because of founder effect speciation, because the ancestor of L. alterna went through a genetic bottleneck when its population crashed at the end of the last ice age, when west Texas turned from woodland into desert.

2. It is an individual L. mexicana that is from a remnant, previously unknown population of L. mexicana in Texas. Even if that is the case, it only proves my hypothesis that L. mexicana once existed in Texas and that it is now absent from most of its original range, including the site where van Devender found vertebral material in packrat middens. My theory is one that is consistent with the fossil record. If anyone has an alternative theory, they still would have to explain how alterna originated and where and how it replaced L. mexicana if the formation of the Chihuahuan Desert was not the cause of such replacement.

3. It is an alterna that has reverted to the ancestral condition in iris color. That possibility, once again, does not affect my theory since it supports my contention that L. mexicana once existed in west Texas and that it has since been replaced by L. alterna. A reversion to an ancestral condition simply proves that L. alterna is derived from L. mexicana, which is my theory.

If none of these scenarios would falsify my hypothesis, then what would? Since my theory is a scientific one, it makes predictions, which are in turn falsifiable. One prediction made by my theory is that genetic variation among the many different populations of L. alterna is low, much lower than the amount of variation among different populations of, say, L. mexicana or L. pyromelana. Hence a finding that L. alterna genetic variation is high would indeed falsify my theory.

Hence I am quite amused to see troy h do the online equivalent of a victory dance (and his comrades dancing along with him) even though his finding that a Davis Mountain L. alterna with a non-silvery gray iris, even if it were authentic, really has no effect on my theory of the origin and evolution of L. alterna."

PS I really would like Troy to explain how the silvery gray iris evolved in L. alterna, if the ancestor of L. alterna did not suffer a population crash and then went through a genetic bottleneck. Tell us how the silvery gray iris is apparently so adaptive that it has replaced the ancestral golden brown iris completely (with but one possible exception) over the entire range of L. alterna.

CKS I really would like Troy to explain how the silvery gray iris evolved in L. alterna, if the ancestor of L. alterna did not suffer a population crash and then went through a genetic bottleneck. Tell us how the silvery gray iris is apparently so adaptive that it has replaced the ancestral golden brown iris completely (with but one possible exception) over the entire range of L. alterna.

Me: Why should I bother to indulge you with a "just-so" story? You asserted that ALL alterna have a silvery gray iris. I demonstrated that you were wrong. End of story. You were wrong, admit it.

BTW, Deserts are defined as areas that receive 10" or less of rainfall per year . . . the Trans-Pecos receives about 18" per year, on average . . . therefore it CANNOT, by definition, be considered a desert (yet another area that you are wrong).

Troy

Troy wrote:
"Why should I bother to indulge you with a "just-so" story? You asserted that ALL alterna have a silvery gray iris. I demonstrated that you were wrong. End of story. You were wrong, admit it."

What we have, finally, is an admission from Troy that he has no explanation of how the silvery gray iris has become a universal trait in Lampropeltis alterna. He has no explanation to account for the replacement of the ancestral golden brown iris by the silvery gray iris in L. alterna. That is because his theory that L. alterna never changed even though its environment changed from woodland to desert since the end of the last ice age cannot explain this fact.

What does one do when one has no explanation for known facts? That is easy for some: dismiss the facts. Troy claims that L. alterna does NOT have a silvery gray iris by showing that one snake, out of the thousands of known specimens, lacks the silvery gray iris. So, he found a photograph of a single snake of unknown origin and claims that this snake proves me wrong because it does not have a silvery gray iris. Even if this specimen is a wild caught L. alterna, and even if it does not have a silvery gray iris, it proves nothing.

I dealt fully with what such a specimen would mean in my previous post, and shows that this single specimen, even if it is authentic, does nothing to change the fact that the silvery gray iris has replaced the golden brown iris and that L. alterna has replaced the ancestral L. mexicana in Texas because west Texas has turned into desert. Troy has no response to my arguments.

Troy's "response" instead is to deny yet another fact. He is claiming once again that West Texas is not a desert. That seems to be something that the animals and plants living in West Texas are unaware of. The plants and animals living there are characteristic of desert environments, and this very simple fact has enabled biologists to call West Texas part of the Chihuahuan Desert. There is even a Chihuahuan Desert Research Institute in West Texas studying the desert biota that exist there. Perhaps Troy should take the people who run this institute to court and make them change the name of this institute to the Chihuahuan Woodland Research Institute, because according to Troy, the Chihuahuan Desert does not exist in Texas.

It is obvious that Troy cannot explain how the silvery gray iris originated in L. alterna, and it is obvious that he cannot explain why L. mexicana no longer lives in Texas or why L. mexicana fails to intergrade with L. alterna. It is obvious that he refuses to accept my theory, which does provide plausible explanations for each of these facts. Therefore Troy has resorted to dismissing the facts that he cannot explain.

The fact that there is no published report of L. alterna without a silvery gray iris calls into question the authenticity of Troy's photograph. If he is willing to back up his claim, he should publish his finding in a scientific journal. I am sure that he will have no problem finding an editor willing to publish it, provided that it is factual piece of information.

You say:

the silvery gray iris has become a universal trait in Lampropeltis alterna

And in the next paragraph acknowledge that some alterna are an exception to this universal rule.

Thank you for finally admitting your error!

Troy

A single exception does not automatically invalidate a rule. I hope that you understand biology well enough to know that the duplication of DNA is not 100% perfect. If it were 100% perfect, then no mutation will ever occur and no evolutionary change will ever occur since there would be no natural variation for natural selection to act upon. The silvery gray iris remains a universal trait in L. alterna even if your claim that you found a wild caught specimen of L. alterna without the silvery gray iris is authentic. A mutation in a single individual does not prove that this trait is not universal. I did not accept your claim as valid. I nevertheless dealt with this find as though it were authentic and see what effect it would have on my theory. As I noted elsewhere, you failed to respond to my arguments, even though my arguments suggest that such a find, even if authentic, does not falsify my hypothesis.

And it remains to be seen whether your claim is true or not. But judging from your persistent refusal to publish your findings, it is perhaps safe to repeat what I have said elsewhere:

There is no beef, only bull, in your claim that you have found an L. alterna without a silvery gray iris.

DHL-- I don't know how long you've been reading this forum, but in case you don't have much experience with CKing (aka CalKing, RSNewton, Lirpa1, etc), you should be aware that he will not change any opinion, will refuse to accept anything done by someone he considers `Hennigian', loves to quote Mayr, and, in general, refuses to accept work done since the 80's. I don't know who he is or what his background is, but he appears to be a troll with a lot of time on his hands and a strong dislike of molecular phylogenetics and modern herpetology.

Patrick Alexander

Thanks for the heads up, but yeah, I should have known better. I've seen his stuff here as well as on fieldherpers and still responded (against better judgement). While I think questioning (even "authorities" is a good idea, arguing to argue is not productive. And one can simply read this thread to see that discussion without comprehension is not productive.

"...arguing to argue is not productive. And one can simply read this thread to see that discussion without comprehension is not productive."

I do not know what you mean by "productive." I made my point. You made yours. It is obvious that we disagree and that we haven't changed each other's minds. It is obvious that many cladists do not give a hoot about heterogeneous taxa. In fact some of them seem to derive some sort of perverse pleasure by deliberately creating heterogeneous taxa even if it can be avoided. That much is clear by their inclusion of Hyla crucifer, Hyla regilla and Hyla cadaverina in Pseudacris. Perhaps they do it deliberately to annoy the Darwinians. Perhaps they are trying to illustrate, by example, what Frost and Etheridge mean by taxa that consist of "my car and 3 geese."

As Moriarty and Cannatella explained, they are merely following "most checklists and field guides" in retaining Hyla regilla, Hyla cadaverina and Hyla crucifer in Pseudacris. That means not all field guides or checklists agree with their taxonomic arrangement. Indeed, the latest edition of R. C. Stebbins' field guide retains Hyla regilla and Hyla cadaverina in Hyla. Stebbins thus shows us that Hennigian classification is by no means universally accepted amongst biologists. In fact, even some cladists are beginning to come to their senses and they are departing from strict Hennigian practices. The recent rejection of Frost and Etheridge's lumping of Chameleonidae and Agamidae, and the resurrection of the family Iguanidae, by some systematists, including Frost and Etheridge themselves, is a good example. Alas, not all cladists have come to their senses. Some of them still insist on straightjacketing themselves with Hennigian dogma.

`Thanks for the heads up, but yeah, I should have known better.'

Unfortunately, it's hard not to get sucked in sometimes. His refined and complex brand of stupid can be hard to ignore...

`I've seen his stuff here as well as on fieldherpers and still responded (against better judgement).'

Ok. I didn't remember seeing you around before, so I wasn't sure.

`While I think questioning (even "authorities" ) is a good idea, arguing to argue is not productive. And one can simply read this thread to see that discussion without comprehension is not productive.'

Seconded, on both counts. The ironic thing, to me, is that I agree with many CKing's basic ideas regarding acceptance of paraphyletic taxa and the like, but I still become a `cladist' when I disagree with his application of the ideas. *shrug*

Patrick Alexander

Patrick wrote:
"Seconded, on both counts. The ironic thing, to me, is that I agree with many CKing's basic ideas regarding acceptance of paraphyletic taxa and the like, but I still become a `cladist' when I disagree with his application of the ideas. *shrug*"

Me:
Unfortunately, that is just another one of Patrick's unsupported assertions. He claims to agree with me, and yet he calls me stupid. It is confounding!

One more piece of advice for Patrick (which will almost certainly be ignored):

Stick to data, not ideology. Cladists will fit the data into their ideology, but scientists will make their theory fit the data. Cladists see the world as being composed of nothing but holophyletic groups. Scientists see that paraphyletic groups are the inevitable result of the process of evolution.

Patrick Alexander wrote:
"he appears to be a troll with a lot of time on his hands and a
strong dislike of molecular phylogenetics and modern herpetology."

Me:
Patrick's claim that I am a "troll" and that I dislike molecular phylogenetics and modern herpetology is so ridiculous that it is actually hilarious. Some of Patrick's responses to my posts in this forum were deleted by the moderator probably because they violated the policy of this forum, since these posts contained asinine remarks and obscenities but no facts or reasoned arguments. He certainly has a lot of chutzpah.

As for the claim that I dislike molecular systematics, that is simply Patrick making yet another unsupported statement. I have not confined my criticism to molecular studies. I have criticized the morphological studies of Kluge and that of Frost and Etheridge, among others. In fact, I used the molecular data from Lopez and Maxson (1995) to show that Old World and New World Elaphe form a monophyletic group and I used the immunological data of Dowling et al. to show that Lampropeltis calligaster is a close relative of Lampropeltis mexicana. In this thread, I have cited Maxson and Wilson's immunological data, which is molecular evidence, to show that it is not a good idea to put Hyla regilla in Pseudacris. I am far from an opponent of "molecular phylogenetics."

Finally, as for the claim that I dislike "modern herpetology." There is indeed a kernel of truth in that statement. Many "modern" herpetologists have straightjacketed themselves with cladistic dogma. As a result, many "modern" systematists are on a rampage to destroy paraphyletic reptilian and amphibian taxa, since cladists (or Hennigians) find such taxa ideologically unacceptable. The cladists' ideological intolerance of paraphyletic taxa and their embrace of such invalid species concepts as the evolutionary species concept and phylogenetic species concept have resulted in a large number of unnecessary taxonomic changes/proposals. The recent transfer of Hyla crucifer, Hyla regilla and Hyla cadaverina to the genus Pseudacris is a very good example. Hedges, who was adhering to cladistic dogma, made the transfer, even though subsequent studies showed that his tree was incorrect and therefore his taxonomic proposal was unnecessary even according to cladistic dogma. If he wasn't adhering to cladistic dogma, his taxonomic proposal would have been unnecessary, saving us all from the taxonomic chaos his proposal has generated. "Modern herpetology" has therefore taken a turn for the worse because of the popularity of cladistic ideology amongst some of its practitioners. Fortunately, there are still many herpetologists who have not succumbed to cladistic dogma. Hence the future of the discipline of herpetology is not entirely hopeless.

Equally fortunately, perhaps, is the fact that Patrick is primarily interested in plants, not reptiles and amphibians. Herpetology's "loss" will be botany's "gain."

Just as the cladists often do when facts are not on their side, Patrick Alexander is choosing to engage in ad hominem. He has no fact to refute any of my arguments concerning the classification or the phylogenetic position of Hyla regilla and he of course does not present any. Yet it is apparent that he does not like my arguments. The facts I presented will not change Patrick's mind at all, even though he has no facts to oppose my arguments. What does Ernst Mayr call people who insist they are right even in the face of opposing evidence? Mayr calls them "dogmatic." Patrick is simply a dogmatic fellow. The minds of the dogmatic cannot be changed, least of all by facts.

Patrick Alexander's unsupported claim that I am someone who somehow "refuses to accept work done since the 80's" is ridiculous to say the least. It is he and many cladists who have often ignored older data because these data contradict either new data or because they contradict cladistic ideology. For example, I accept the mtDNA data of Lopez and Maxson (1995), which shows that Old World and New World Elaphe together form a monophyletic group. Patrick does not, because he thinks that Elaphe is polyphyletic. I have no problem with Rodriguez-Robles et al.'s data on the phylogeography of Lampropeltis zonata, and I have no problem with Rodriguez-Robles' mtDNA data on Charina bottae, even though I disagree with their conclusion that the rubber boa should be divided into two species. The mtDNA data of L. zonata and C. bottae by Rodriguez-Robles et al. are published within the last few years, thus demonstrating that Patrick is simply incorrect in his delusional assertion. Unlike Patrick, I do not judge data by their date. Unlike milk, data has no expiration date.

My advice to Patrick is this: you are entitled to your opinion but you should support your opinion with facts, not fiction.

CKing said "All alterna have silvery gray irises"

I refuted that assertion (quite easily) by going into my animal room and shooting a photo of one of my wild-caught Davis Mt alterna (see photo above).

CKing went away from the forums for several days, and when he returned, he stammered "Uh, its either not an alterna, or a mutant, or a reversal to a more primitive form".

A more principled person would have simply acknowledged THAT HE WAS WRONG!

Oh, and since the Trans-Pecos region of west Texas receives more than 10" of rainfall on average per year, it cannot be precisely described as a true Desert - since, by definition, deserts receive 10" of rainfall or less, on average, per year.

Troy

Facts do matter. That is why I do not ignore immunological data, unlike Troy.

Troy has kindly provided us with a nice photograph of one of his animals. He claims that it is a wild caught L. alterna from the Davis Mountains. If so, perhaps he should publish this piece of evidence, since facts are indeed very important.

That said, I am willing to give Troy the benefit of the doubt and I have discussed at length how such a discovery will affect my theory that L. mexicana became extinct and became replaced by L. alterna. Rather than addressing my arguments, Troy has chosen first to ignore them and now he has misrepresented them. I will repost the arguments I made below and let the readers judge for themselves whether Troy has answered the points I made.

I will not, for the time being, contest troy h's claim that someone has collected an alterna without a silvery gray iris. I will instead try to figure out what such a find, if it were authentic, would do to my theory of how L. mexicana transformed into L. alterna.

Of course, if it were a fake, then it would not change my theory. Assuming, however, that it is real, I will see what effects it will have on my theory.

1. It is a mutant. Mutations happen sooner or later. If it is a mutant then it has no effect on my theory. In fact, it supports my contention that the silvery gray iris (or iris color in general) in L. alterna is a selectively neutral character that became fixed because of founder effect speciation, because the ancestor of L. alterna went through a genetic bottleneck when its population crashed at the end of the last ice age, when west Texas turned from woodland into desert.

2. It is an individual L. mexicana that is from a remnant, previously unknown population of L. mexicana in Texas. Even if that is the case, it only proves my hypothesis that L. mexicana once existed in Texas and that it is now absent from most of its original range, including the site where van Devender found vertebral material in packrat middens. My theory is one that is consistent with the fossil record. If anyone has an alternative theory, they still would have to explain how alterna originated and where and how it replaced L. mexicana if the formation of the Chihuahuan Desert was not the cause of such replacement.

3. It is an alterna that has reverted to the ancestral condition in iris color. That possibility, once again, does not affect my theory since it supports my contention that L. mexicana once existed in west Texas and that it has since been replaced by L. alterna. A reversion to an ancestral condition simply proves that L. alterna is derived from L. mexicana, which is my theory.

If none of these scenarios would falsify my hypothesis, then what would? Since my theory is a scientific one, it makes predictions, which are in turn falsifiable. One prediction made by my theory is that genetic variation among the many different populations of L. alterna is low, much lower than the amount of variation among different populations of, say, L. mexicana or L. pyromelana. Hence a finding that L. alterna genetic variation is high would indeed falsify my theory.

Hence I am quite amused to see troy h do the online equivalent of a victory dance (and his comrades dancing along with him) even though his finding that a Davis Mountain L. alterna with a non-silvery gray iris, even if it were authentic, really has no effect on my theory of the origin and evolution of L. alterna.

PS I really would like Troy to explain how the silvery gray iris evolved in L. alterna, if the ancestor of L. alterna did not suffer a population crash and then went through a genetic bottleneck. Tell us how the silvery gray iris is apparently so adaptive that it has replaced the ancestral golden brown iris
completely (with but one possible exception) over the entire range of L. alterna.

Troy wrote:

"A more principled person would have simply acknowledged THAT HE WAS WRONG!

Oh, and since the Trans-Pecos region of west Texas receives more than 10" of rainfall on average per year, it cannot be precisely described as a true Desert - since, by definition, deserts receive 10" of rainfall or less, on average, per year."

Me:
According to Troy's definition of a desert and according to his own claim, Val Verde County is not a desert. Below is a photograph of Val Verde County near Langtry. The treeless, rocky area within this photograph is NOT a desert according to Troy! And of course I am wrong for claiming that the Chihuahuan Desert is a desert according to Troy! Since there are a few tufts of grass at the side of the road, perhaps we should call this area a grassland instead. Or may be we should call this a woodland habitat because, according to Troy, L. alterna lives in woodland habitats but it is not a snake adapted to the desert. Since the area in this photograph is prime L. alterna habitat, and since L. alterna is not a desert reptile, we can never call this area a desert, especially since it receives more than 10 inches of rainfall per year according to Troy.