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The Center for North American Herpetology
Lawrence, Kansas
http://www.cnah.org
8 April 2007

HOW AND WHEN DID OLD WORLD RAT SNAKES DISPERSE INTO THE NEW WORLD?

Frank T. Burbrink and Robin Lawson
2007. Molecular Phylogenetics and Evolution 43: 173-189

Abstract: To examine Holarctic snake dispersal, we inferred a phylogenetic tree
from four mtDNA genes and one scnDNA gene for most species of the Old World
(OW) and New World (NW) colubrid group known as rat snakes. Ancestral area
distributions are estimated for various clades using divergence–vicariance
analysis and maximum likelihood on trees produced using Bayesian inference.
Dates of divergence for the same clades are estimated using penalized likelihood
with statistically crosschecked calibration references obtained from the Miocene
fossil record. With ancestral areas and associated dates estimated, various
hypotheses concerning the age and environment associated with the origin of rat
snakes and the dispersal of NW taxa from OW ancestors were tested. Results
suggest that the rat snakes originated in tropical Asia in the late Eocene and
subsequently dispersed to the Western and Eastern Palearctic by the early
Oligocene. These analyses also suggest that the monophyletic NW rat snakes
(the Lampropeltini) diverged from OW rat snakes and dispersed through Beringia
in the late Oligocene/early Miocene when this land bridge was mostly composed
of deciduous and coniferous forests.

*****

Available as a download from the CNAH PDF Library at

http://www.cnah.org/cnah_pdf.asp

*****

CNAH Note: The title will mislead you. The most significant information in this
very thorough and significant paper is the proposal that the genus Pantherophis
(New World Rat Snakes formerly placed in the genus Elaphe) should be placed in
the synonymy of the genus Pituophis (Bullsnakes, Gopher Snakes, and Pine
Snakes). This change is formally suggested in the last sentence of the last
paragraph of the paper.

The resulting scientific taxonomy (and correct spelling of the specific names)
for the nine North American colubrid species affected would be:

Eastern Rat Snake (Pituophis alleghaniensis)
Baird's Rat Snake (Pituophis bairdi)
Great Plains Rat Snake (Pituophis emoryi)
Eastern Fox Snake (Pituophis gloydi)
Eastern Corn Snake (Pituophis guttatus)
Western Rat Snake (Pituophis obsoletus)
Slowinski's Corn Snake (Pituophis slowinskii)
Midland Rat Snake (Pituophis spiloides)
Western Fox Snake (Pituophis vulpinus)

Standard common names would remain the same; the importance of using such
standardized common names looms large in the face of such a significant and
far-reaching change as proposed by Burbrink and Lawson.

I made a similar one in cornsnakes that says:

For those that jumped on the Pantherophis bandwagon too soon, ready to kiss it bye? The new paper by Burbrink Lawson (2007) just groups all New World Lampropeltini into "Pituophis." Of course, the paper is by Burbrink who has a phenomenal record for NEVER getting anything correct, so take it at face value. Those that jump on the "new hypothesis" band wagon before confirmation with further scientific studies make me chuckle.....

Humorously yours. By the way, the publication is real.
KJ
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KJUN Snakehaven

>>For those that jumped on the Pantherophis bandwagon too soon, ready to kiss it bye?

Not that I have any authority on the subject (because I don't), but I work for a couple of PhD level herpetologists who specialize in the systematics of herps.
My job is to recurate and update accepted taxanomic revisions in the museum's herp collection. Well, being a ratsnake addict, I frequently ask Dr. Jack Sites and Dr. Dan Mulcahy stuff about ratsnake systematics. One of those questions that I have asked is, "Why haven't we changed our North American Elaphe specimens to Pantherophis? I've asked them both this, and they have each told me that the proposal has very little support in the professional herp systematics community. It seems that the only group that has accepted this change is the reptile pet trade. I know that this may be construed as blasphemy in some circles , but the herpetocultural and pet trade communities are NOT scientific communities, collectively speaking.

It's obvious that there needs to be a taxonomic revision between NW and OW "ratsnakes", but we lay herpers first need to look at what is accepted by those whose business it is to make and critique taxanomic revision proposals, and not be eager to make a change based on one largely unsupported paper, which seems to be the case with Pantherophis.

And though the CNAH is frequently referenced in the herp community, the association is viewed as controversial, at best, within the doctorate-level systematist community. The CNAH does not have near the support as say, the SSAR or the EMBL database, and many professional herpetologists question the validity of CNAH.

DR
Suboc.com

I honestly didn't have time to read all the post, but if you ask them WHY they'd probably add some info about the "10 year rule" or something like it. MANY of them don't make changes for at least 10 years to see if the hypothesis is supported, rejected, or questioned further. Heck, the Texas Cooperative Wildlife Collections still calls E. emoryi by the old name (E. guttata) in their records. Why? Because name changes are hypotheses and NOTHING more. Hobbyists talk bad about how often names change, but that is because they don't realize that a publication is a hypothesis and NOT a factual name change.

The Pantherophis bandwagon has just cracked me up because I knew they'd just get mad over the "next change" and wonder why they couldn't leave things alone. Nobody would listen to me (hobbyists), so I'm just curious to see which ones start calling them Pituophis so quickly OR are hypocritical and stick with Pantherophis. I guess we'll see, huh? LOL.

In my recent articles on Kisatchie cornsnakes, I explain why I still use Elaphe. It is because there isn't acceptance in the SCIENTIFIC community yet. My hobby is with laymen and I love this culture and it's participants, but my job as a herpetologist takes precedence. It is still Elaphe to me, even though I think the NW and OW split is warranted. This new publication seems to have some unspoken assumptions that aren't (yet) supported by fossil records that would be required fore me to accept this as-is.

Ever notice that NOTHING of Burbrink's makes much biological sense and is poorly accepted by the scientific community outside of his clique. His black ratsnake division is based on the assumption that ratsnakes can't swim across rivers.....lol.

Thanks for your post,
KJ
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KJUN Snakehaven

>>I honestly didn't have time to read all the post, but if you ask them WHY they'd probably add some info about the "10 year rule" or something like it. MANY of them don't make changes for at least 10 years to see if the hypothesis is supported, rejected, or questioned further. Heck, the Texas Cooperative Wildlife Collections still calls E. emoryi by the old name (E. guttata) in their records. Why? Because name changes are hypotheses and NOTHING more. Hobbyists talk bad about how often names change, but that is because they don't realize that a publication is a hypothesis and NOT a factual name change.
>>
>>The Pantherophis bandwagon has just cracked me up because I knew they'd just get mad over the "next change" and wonder why they couldn't leave things alone. Nobody would listen to me (hobbyists), so I'm just curious to see which ones start calling them Pituophis so quickly OR are hypocritical and stick with Pantherophis. I guess we'll see, huh? LOL.
>>
>>In my recent articles on Kisatchie cornsnakes, I explain why I still use Elaphe. It is because there isn't acceptance in the SCIENTIFIC community yet. My hobby is with laymen and I love this culture and it's participants, but my job as a herpetologist takes precedence. It is still Elaphe to me, even though I think the NW and OW split is warranted. This new publication seems to have some unspoken assumptions that aren't (yet) supported by fossil records that would be required fore me to accept this as-is.
>>
>>Ever notice that NOTHING of Burbrink's makes much biological sense and is poorly accepted by the scientific community outside of his clique. His black ratsnake division is based on the assumption that ratsnakes can't swim across rivers.....lol.
>>
>>Thanks for your post,
>>KJ
>>-----
>> KJUN Snakehaven

"Heck, the Texas Cooperative Wildlife Collections still calls E. emoryi by the old name (E. guttata) in their records. Why? Because name changes are hypotheses and NOTHING more."

curious..
why do you feel that corn/emoryi crosses are hybrids when for years and years emoryi was listed as a sub species to guttata (elaphe g. emoryi) especially when in your own words....name changes are hypotheses and NOTHING more?
what makes THAT change more then hypotheses?
-----
Corn snakes and rat snakes..No one can have just one.
"Resistance is futile"
Jimmy Johnson
(Draybar)
Draybars Snakes

_____

>>why do you feel that corn/emoryi crosses are hybrids when for years and years emoryi was listed as a sub species to guttata (elaphe g. emoryi) especially when in your own words....name changes are hypotheses and NOTHING more?
>>what makes THAT change more then hypotheses?

Those aren't my words - I just repeat what they are - by definition (in reference to "hypothesis".

The first of the RECENT splits of Elaphe emoryi versus Elaphe guttata was proposed in the '70's. Very little refuted that, and it is only beginning to get acceptance. Time. That's why I accept it. It has pretty much stood the test of time and additional studies, with some minor exceptions. All are available with a decent literature search, so I'm sure you've seen them if you care.

Finally, intergrades, by definition, are susbecpific crosses IN NATURE. Hybrid is a better definition for man-made subspecific crosses even though hobbyists use the term "intergrade" erronously to describe subspecific crosses done in captivity. Intergrade as a VERY specific definition in these fields. Hybrid can mean everything from crosses among different species to crosses within a species of different lines ("hybrid" mice in research are all the same species, for example). Intergrade is an exclusive term - hybrid is an inclusive term.

Frequently, you'll see me use the term "hybrid" for corsses between or among species, intergrades for natural subspecific crosses, and subspecific hybrid to descibed man-made crosses between or among different subspecies.

KJ
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KJUN Snakehaven

>>>>why do you feel that corn/emoryi crosses are hybrids when for years and years emoryi was listed as a sub species to guttata (elaphe g. emoryi) especially when in your own words....name changes are hypotheses and NOTHING more?
>>>>what makes THAT change more then hypotheses?
>>
>>Those aren't my words - I just repeat what they are - by definition (in reference to "hypothesis".
>>
>>The first of the RECENT splits of Elaphe emoryi versus Elaphe guttata was proposed in the '70's. Very little refuted that, and it is only beginning to get acceptance. Time. That's why I accept it. It has pretty much stood the test of time and additional studies, with some minor exceptions. All are available with a decent literature search, so I'm sure you've seen them if you care.
>>
>>Finally, intergrades, by definition, are susbecpific crosses IN NATURE. Hybrid is a better definition for man-made subspecific crosses even though hobbyists use the term "intergrade" erronously to describe subspecific crosses done in captivity. Intergrade as a VERY specific definition in these fields. Hybrid can mean everything from crosses among different species to crosses within a species of different lines ("hybrid" mice in research are all the same species, for example). Intergrade is an exclusive term - hybrid is an inclusive term.
>>
>>Frequently, you'll see me use the term "hybrid" for corsses between or among species, intergrades for natural subspecific crosses, and subspecific hybrid to descibed man-made crosses between or among different subspecies.
>>
>>KJ
>>-----
>> KJUN Snakehaven

thanks..
Actually you did not reference those words to anyone else so one could only take them to be YOUR words. Easy mistake.
I have a few books that were published in the 90's that still have them listed as elaphe g. emoryi
so I guess it just depends on WHERE you do the research...huh?
-----
Corn snakes and rat snakes..No one can have just one.
"Resistance is futile"
Jimmy Johnson
(Draybar)
Draybars Snakes

_____

>>Actually you did not reference those words to anyone else so one could only take them to be YOUR words. Easy mistake.

When you say the sky is blue, do you reference the definition of sky or blue? That's the case here. People may not understand it, but all systematic "conclusions" are hypotheses of relationships. Nothing more.

>>I have a few books that were published in the 90's that still have them listed as elaphe g. emoryi
>>so I guess it just depends on WHERE you do the research...huh?

LOL, nah. That's not it at all. It depends on the books. Many hobbyists confuse the difference between information in peer-reviewed sources and popular books/magazines.
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KJUN Snakehaven

I am not an academic and will not claim to be, but I do have some 'opinions' about this. It seems obvious that our NW rats are different enough from OW (Elaphe, etc.) to warrant being placed in a separate genus, whatever it may be. However, they still seem like they are different enough from Pituophis to remain in a separate genus from them. I am not that attached to taxonomic names and I understand that they represent hypotheses that are often not accepted by the majority of the scientific community. The proposal of switching the North American Pantherophis/Elaphe to Pituophis doesn't bother me at all, and with all of the stuff that Burbrink comes up with, it is to be expected. I don't agree with most of what I have seen from Burbrink either. I don't really care what the genus is except that it is convenient to differentiate animals, and sticking with Pantherophis is as good as sticking with Elaphe, imho. FWIW, Elaphe snakes were not always consistently called Elaphe either, eg: 'Elaphe' obsoleta obsoleta was called Coluber obsoletus in 1823, Scotophis alleghaniensis in 1853 (Baird & Girard) and the 'hypothesized' name Pantherophis is not new either and was suggested by Garman in 1892 (information taken from "A Monograph of the Colubrid Snakes of the Genus Elaphe Fitzinger" 1996 by Klaus-Dieter Schulz). I don't really feel that it would be hypocritical to continue to use the name Pantherophis rather than jumping on the Pituophis bandwagon, and would it be less hypocritical to go back to using Elaphe? I sometimes use Pantherophis and Elaphe interchangeably (Pantherophis/Elaphe) in an effort to placate both sides. I am more interested in the animals, themselves, than what we call them.

-Toby Brock

>>FWIW, Elaphe snakes were not always consistently called Elaphe either, eg: 'Elaphe' obsoleta obsoleta was called Coluber obsoletus.

Well, actually almost every snake that isn't venomous was in the genus Coluber at one time or another, including all NA Elaphe and Bogertophis. Probably Senticolis too.
Even the African rock python and Indian python were once called Coluber sebae and Coluber molurus, respectively.

DR

Suboc.com

Dusty, I actually knew that also, and just used it as an example of the changeable nature of taxonomic names. Elaphe obsoleta also went under the genera; Scotophis, Elaphis and Pantherophis.

-Toby

>>Dusty, I actually knew that also, and just used it as an example of the changeable nature of taxonomic names.

Cool. I was sure that you did know. I just felt like posting that for all the readers.

DR

Don't forget that Pituophis was once Pityophis, too.

Lampropeltis was once Ophidiobolus (or was it Ophidiabolus? - I think it "o" is correct...). I love that name. The root words EXACTLY describe the diet of kingsnakes....lol.
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KJUN Snakehaven

In essence, all they did was back up a node and say "THIS" node should be the genus and all branches off of it should be different species. Made the genus more inclusive (and general). The designations of groups from species on up is relatively arbitrary, and different people can look at the same tree and come to different conclusions on how to group them. That's really minor.

KJ
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KJUN Snakehaven

Thanks KJ,

Like I said, I am not an academic, just a hobbyist, but the Pantherophis hypothesis seemed like a step forward, being more exclusive than the old Elaphe taxon, while the proposed change to Pituophis seems like a big step backward, being more inclusive.

-Toby

Thanks KJ.

I will continue to use Elaphe until Pantherophis is made official.

Lumping E. guttata with Pitophis is patently absurd.

I have a big problem with the cladistic/statistic/mtDNA approach to taxonomic classification. The problem I have is it contradicts observable physical evidence. Perhaps this is unscientific, but any competent observer looking at a corn, black rat, fox and pine will see that the first three are more like each other than any of them are like the last. Furthermore, I feel that the taxo-babble jargon used to justify this wholesale renaming is effectively an indecipherable smoke screen misleading laypeople into jumping on the bandwagon. This may not be deliberate, but it certainly is effective.

If Burbink and the rest of the taxonomic "Drive-Bys" have their way, we may as well trash all our copies of Conant, Tenant, and what-have-you field guides because they are all useless. We are fast approaching the point where nothing short of a pocket DNA analyzer will be required to determine what anything is in the field.

Regards,

John DeMelas
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I am so not lesdysxic!

0.1 Creamsicle Cornsake "Yolanda"
1.0 Bairds Ratsnake "Steely Dan"
0.1 Desert Kingsnake "FATTY"
0.1 Black Rat "Roberta" RELEASED!!!

..I should have read the others..
:Mark
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Surrender Dorothy!

www.crimsonking.funtigo.com

the firestorm of dissenting opinions on the changes in taxonomic designations, three items caught my attention.

One was that Coronella is identified as the closest living relative to all the new world ratsnakes and related genera (lampropeltines). I find that pretty interesting.

The other is that the fox snakes are the sister clade to the pits (pines, bulls and gophers). That makes them closer to pits than to obsoleta or guttata. The reason I find that interesting is several years ago Bart Bruno stated that he believed the fox snakes were pits masquerading as ratsnakes. He said that they even had similar musk to each other. Looks like he may be right.

Third is that schrencki is the closest relative to the european four lined ratsnake. Very interesting.

Regards
John

>>The other is that the fox snakes are the sister clade to the pits (pines, bulls and gophers). That makes them closer to pits than to obsoleta or guttata. The reason I find that interesting is several years ago Bart Bruno stated that he believed the fox snakes were pits masquerading as ratsnakes. He said that they even had similar musk to each other. Looks like he may be right.

That was proposed atleast 30 years ago - probably longer if we cared to look. It isn't an original idea. What is LIKELY, but I believe unproved by the fossil record as of today, is that vulpina/gloydi are closer to the ancestral form that gave rise to NA ratsnakes and Pituophis than either of those two are to the ancestor.
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KJUN Snakehaven

for that idea about the fox/pit connection? I'd be interested to see what it was based on.

Regards
John

>>for that idea about the fox/pit connection? I'd be interested to see what it was based on.

Talk to Doug Rossman if you can find contact info for him. He's the one that put the idea in my head back when I was a wee college student under his instruction.........lol. I, personally, can't think of a reference off of the top of my head, but I know I have read it since in some of theolder books. I don't know if it was Shaw, West, Wright, or..... ??? It's been a long time - sorry.

ALTHOUGH, if you look at how they get allied in just about all taxonomic trees, it is self-evident. Vulpina/Gloydi would likely make an excellent outgroup for future Pituophis studies. I'm referring, of course, to the current Pituophis genus and not the proposed Pituophis genus.
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KJUN Snakehaven

>>Talk to Doug Rossman if you can find contact info for him. He's the one that put the idea in my head back when I was a wee college student under his instruction.........lol. I, personally, can't think of a reference off of the top of my head, but I know I have read it since in some of theolder books. I don't know if it was Shaw, West, Wright, or..... ??? It's been a long time - sorry.

Could it have been Dowling's paper (from the 70s?) that you are thinking of? Not sure of the fox/pit connection but I believe his work suggested that e. obsoleta was more closely related to p. melanoleucus than to e. guttata, IIRC. (It's been awhile though, I could be way off on this.)

dg

I'd like to correspond with him too. It's good to know that there's at least one other person on the planet that sees a connection with the Fox Snakes and Pituophis. I wonder if he took as much ridicule as I did for espousing such a view?

Bart

I personally don't think this is going to be accepted.

I also don't agree with this taxonomic interpretation of his tree. I think he throws the baby out with the bathwater.

The tree is interesting for a couple of reasons -

1. It does show the close relationship between pits and foxsnakes.

2. It shows how distant Bogertophis is from the rest of the NA rats (they are closer to kingsnake/glossy/scarletsnake clade).

3. It shows that guttata shouldn't be in the same genus as obsoleta and bairdii.

Here's another less problematic approach to his tree...

- Pantherophis guttata
- Pantherophis slowinskii
- Pantherophis emoryi

Because Pantherophis only includes the cornsnake group, the real NA ratsnakes (obsoleta and bairdii - let's ignore alleghaniensis, etc for now) can't be in that genus. They have to be in their own genus, but they can't be Elaphe either. So I believe the oldest valid name for the group would be the genus Georgia.

- Georgia obsoleta
- Georgia bairdii

- Pituophis melanoleucus
- P. vulpinus
- P. gloydi
- P. catenifer
- P. sayi
- P. deppei
- P. lineaticollis

- Bogertophis stay as they are now

This is still more evidence of how paraphyletic/polyphyletic NA ratsnakes were.
-----
Chris Harrison
San Antonio, Texas

This resolution makes very good sense to me, Chris. Thanks for posting it!

-Toby Brock

I've been working with gloydi and vulpina since 1999. They are extremely interesting (and finicky). I've always wanted to see the fox snakes studied. I'm sure it'll take another ten years for the taxonomists to come to a consensus but I'm pleased that a number of DNA studies are being done on the species and genera that I've always been interested in.

But the fact that Utiger and Burbrink have come up with completely different locations for prasina and frenata leads me to believe that somebody is mistaken. Time will tell (along with more data).

Regards
John

im no taxanomical expert, but putting ratsnakes in the same genus as bulls/pines seems a bit ridiculous to me? i mean im sure there is a lot of dna evidence and whatnot that i dont know about, but, personally i think those snakes look totally different. and i know taxonomy is more than looks but look at a rat snakes head then look at a pine snakes head, i think pituophis are more closely related to lizards than ratsnakes to tell you the truth.

>>im no taxanomical expert, but putting ratsnakes in the same genus as bulls/pines seems a bit ridiculous to me? i mean im sure there is a lot of dna evidence and whatnot that i dont know about, but, personally i think those snakes look totally different. and i know taxonomy is more than looks but look at a rat snakes head then look at a pine snakes head, i think pituophis are more closely related to lizards than ratsnakes to tell you the truth.

I'm taking the lizard comment as a joke, but don't forget that head shape in ratsnakes is due to phenotypic plasticity and not just genetics, so don't put much weight on that characteristic.

Looks don't mean much - they are often misleading when it comes to how organisms are related to each other. I don't like this new hypothesis, but the logic IS there. I just disagree with his conclusions. Of course, Burbrink hasn't gotten anything correct yet.....
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KJUN Snakehaven

This may seem like an ignorant question but.....

"3. It shows that guttata shouldn't be in the same genus as obsoleta and bairdii."

Why do you believe that corns should be seperated from the other ratsnakes?
-----
.1 Snow Corn "Hope"
1. Orange Albino Black Ratsnake "Chunk" (Goonies)
.1 Orange Albino Black Ratsnake "Peaches"
.1 MO Locale Black Ratsnake "Molly" (Flogging Molly)

"Have you ever tried simply turning off the T.V., sitting down with your kids... and hitting them?"

See Figure 3 in the article. Elaphe guttata is the oldest lineage from that group. THAT node must include all species descended from it to be monophyletic. Burbrink calls in a genus. I still consider it larger group than a genus. He DOES make in a monophyletic group his way, though.

OK, notice that the E. guttata lineage is the oldest one. It breaks off before the E. obsoleta/bairdi group. If that tree is correct, and you put E. guttata with E. obsoleta/bairdi, then you HAVE to group all of them in the same genus as P. melanoleucus if you want it to me monophyletic. SO, again IF that tree is correct, the other option would be to have corns in their OWN genus.

Of course, this tree might not be correct. Science never knows TRUTH - this is just a hypothesis based on the data to be further tested.

KJ

>>This may seem like an ignorant question but.....
>>
>>"3. It shows that guttata shouldn't be in the same genus as obsoleta and bairdii."
>>
>>Why do you believe that corns should be seperated from the other ratsnakes?
>>-----
>>.1 Snow Corn "Hope"
>> 1. Orange Albino Black Ratsnake "Chunk" (Goonies)
>>.1 Orange Albino Black Ratsnake "Peaches"
>>.1 MO Locale Black Ratsnake "Molly" (Flogging Molly)
>>
>>"Have you ever tried simply turning off the T.V., sitting down with your kids... and hitting them?"
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KJUN Snakehaven

Here's a lower impact scenario:

Why not transfer all Pituophis species to the Pantherophis genus instead. The result would be a nice and clean monophyletic taxon, which is just as plausible, perhaps even more parsinomiously, than three smaller monophyletic taxa.

Very interesting article, regardless wether their conclusions get accepted or not.

>>Why not transfer all Pituophis species to the Pantherophis genus instead. The result would be a nice and clean monophyletic taxon, which is just as plausible, perhaps even more parsinomiously, than three smaller monophyletic taxa.

Because the use of the genus Pituophis predates the use of the genus Pantherophis. Oldest one wins. If it gets combined, they it has to be Pituophis.

KJ
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KJUN Snakehaven

The oldest name prevails when it concerns the same taxon. In this case we're talking about a new and different taxon, N.A. Ratsnakes including Pine, Bull and Gopher Snakes that is. In the cladograms as proposed by Burbrink & Lawson, Pantherophis is basal to Pituophis. Therefore it can be argumented that Pituophis should be synonimized with Pantherophis, instead of applying the oldest name rule.

Interesting note Chris. Here's your approach:

- Pantherophis guttata
- Pantherophis slowinskii
- Pantherophis emoryi

- Georgia obsoleta
- Georgia bairdii

- Pituophis melanoleucus
- P. vulpinus
- P. gloydi
- P. catenifer
- P. sayi
- P. deppei
- P. lineaticollis

- Bogertophis stay as they are now

My own approach, based on nothing more than gut feeling, would go like this:

-Pantherophis guttatus ssp.

-Georgia obsoleta ssp.
-Georgia bairdi

-Scotophis vulpinus
-Scotophis gloydi

-Pituophis melanoleucus ssp.
-Pituophis catenifer ssp.
-Pituophis sayi ssp.
-Pituophis ruthveni (sayi ruthveni?)
-Pituophis deppei (ssp.?)
-Pituophis lineaticollis (ssp.?)

-Bogertophis

Basically, I think that the fox snakes deserve generic status. Splitting emoryi from guttatus would be easy if it weren't for the "slowinskii" snakes from LA, TX, AR, and OK......

Shane

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Lifelist