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Pales and Reds - thoughts and reflection

Sunherp Feb 17, 2009 11:08 AM

Sorry for such a long post...

A thread below (by Doug L.) has spurred a little discussion regarding the range of syspila and mulitstrata in Nebraska. First, let me state that the monograph published by Ken Williams on the species Lampropeltis triangulum is an outstanding work. It’s the best thing we’ve got to work with so far, but it’s not a pinnacle work. There’s always room for improvement as further info is uncovered.

It is well documented that the form of Lampropeltis triangulum known as mulitstrata occurs in the eastern 2/3 Montana, western South Dakota (WEST of the Missouri River), parts of Wyoming, and northern Nebraska. This form intergrades (experiences mass gene flow) with the form known as gentilis to the south in northern Colorado and southern Nebraska. There is considerable debate as to whether mulitstrata and gentilis should be considered distinct from one another, as there are few, if any, discrete characteristics to distinguish one from the other (physically or biochemically). We’ll leave this little wrinkle out for this discussion (unless someone really wants to discuss it…LOL).

The mulitstrata form is known, as stated above, to be restricted west of the Missouri River in SD, while the form we know as syspila is found in a few counties to the east of the Missouri River within that state. The animals from along the river valley display characteristics of both forms. Why is this? Well, obviously there’s gene flow between the two populations. The river is no barrier to milksnake movement, as Dell can (will?) attest. Also, anyone who’s ever been to that area knows that the Missouri River in the Dakotas and Nebraska is sort of a demarcation between a more arid, xeric habitat (mulitstrata habitat) and a wetter, more mesic habitat (syspila habitat). One way to look at the situation is that the river valley provides a transition habitat, where one phenotype isn’t preferred over another. This area of non-differentiation is called a “primary intergrade zone” by those who study this sort of thing.

(I think wikipedia has a good, solid explanation of the different types of intergrades – “temporalis” along the eastern sea board is more like what we call a “secondary intergrade” with some possible reticulate introgression).

How does this translate into what we’re seeing in Nebraska with Doug’s animals? Williams, in his 1988 edition of his milksnake monograph, described animals from extreme north-eastern Nebraska, along the Missouri River and the South Dakota state line, as being mulitstrata X syspila intergrades. He lacked specimens along the Missouri River to the south of these, but as new specimens are turned up from this area (as Doug’s were), the picture becomes more complete. Intergradient animals are found all along the eastern portion of Nebraska, along the Missouri River Valley, and south (in the form of syspila X gentilis intergrades) along the Flint Hills of Kansas. The Flint Hills of KS provide a similar habitat transition zone as the Missouri River Valley does to the north.

How do we know the animals Doug posted are intergrades? Well, aside from locality, we can see that the animals lack ventro-lateral blotches, a requisite of syspila and Lampropeltis triangulum triangulum. Secondly, we look at their head pattern. Some of the animals show a nice red cap, as a “good” syspila should. Others, from the same locality, show a black or mottled head cap, as a “good” gentilis or mulitstrata should. The very fact that the region produces animals which, as a group, are unidentifiable to subspecies evidences (strongly) that it’s an intergrade zone.

Please, add your thoughts, comments, criticisms, and photos!

-Cole Grover

Thomas Co., NE

Pennington Co., SD

Image

Replies (54)

shannon brown Feb 17, 2009 11:18 AM

Great Post man, you knocked it out of the park right there man.Without locality most of these animals are just North American Milks.

L8r
Image

Sunherp Feb 17, 2009 11:21 AM

That's a looker, Shannon! Feel free to send it up here when you get tired of it... LOL

Thanks for the compliments. I've been thinking about a post like this for quite a while, and the discussion below made it feel like the right time.

-Cole

Yellowstone Co., MT ventral shot
Image

snake_bit Feb 17, 2009 01:09 PM

Nice work Cole,
I expect you to do the pinnacle work on this subject one day.
.
Seems that extreme eastern KS milks can have dorsal lateral blotches, no dorsal lateral blotches or a mix of both. Does that make eastern KS a intergrade area? Peterson calls that a syspila zone but the snakes don't read those books.Here are two milks from the same hillside in east KS.(two counties east of MO).
Note that one has dorsal lateral markings and the other does not.


You remember this one from deep in gentilis range.

Looks more like a pale then many pales I see yet its not in the correct area to be a pale.

I think we have all seen red heads fron lincoln russell and ellis but thay are rare.

Now here is a smokin intergrade fron KS flint hills.
This guy ROCKS






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"Wake me when its April"

Doug L

Sunherp Feb 17, 2009 01:23 PM

Doug,

I think in Eastern Kansas we're seeing some physical signs of gene flow from the adjacent Flint Hills region. That animal (Flint Hills) is stunning, no doubt.

snake_bit Feb 17, 2009 01:42 PM

Cole Do you have pics of MO milks? I would like to see a dozen or two from western MO.Maybe ill do a search on the fieldherpers forum.I have only found one from MO and it was from way east.I just wonder if any of those have zero dorsolaterl markings.
Its my believe that these are all the same snake. talori gentilis multistrata,syspilia and celaenops.I say throw them all in the same bin and call it regional variation.
BTW Do you have a Williams range map?


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"Wake me when its April"

Doug L

Sunherp Feb 17, 2009 02:13 PM

Doug,

I do have the Williams range map, but it's in the book and I don't have a scanner. It shows northeastern NE as a definite intergrade zone, and a lack of specimens from the east-central part of that state.

I think the most recent genetic studies found that syspila clustered with triangulum, while gentilis, taylori, and multistrata all clustered together and close to (with?) celaenops, amaura, and annulata. That arrangement would leave us with 3 (or 2) North American subspecies, not counting the enigmatic elapsoides.

Ventro-lateral blotches may not be in ever example of "pure" syspila, but they're pretty damn common. Their absence may be a sign of gene flow from more "banded" animals (gentilis or multistrata, for example).

Here's a few animals from extreme western MO. Not my photos - they were taken by Matt Jepson and stolen by me.

-Cole


Image

snake_bit Feb 17, 2009 02:42 PM

Thanks for this post Cole I think it clears some things up in my mind.
Macro pic that Williams map when you get home and try to post it.
Mat Jepson has some nice critters,love those two bright orange and thick black ones


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"Wake me when its April"

Doug L

terryd Feb 17, 2009 04:48 PM

Really interesting stuff guys. Wish I had more time to respond.
But I did shoot these off quick.

Systematics & Natural History of the American Milk Snake,
Lampropeltis triangulum

Range map

I also like this book.
Kingsnakes & Milk snakes, by Ronald G. Markel.

Okay I gotta go, busy, busy, today.
-Dell

Sunherp Feb 17, 2009 05:10 PM

I knew you'd come through! LOL

-Cole

snake_bit Feb 17, 2009 06:04 PM

Thanks Dell Ill save these
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"Wake me when its April"

Doug L

Dniles Feb 18, 2009 06:11 AM

Hey Dell,

thanks for posting these maps. very cool. I just picked up Markel's book at a herp show last weekend. I need to get my hands on Systematics & Natural History of the American Milk Snake for sure. A whole book dedicated to triangulum - what could be better??

Dave
DNS Reptiles

Sunherp Feb 18, 2009 10:48 AM

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Dniles Feb 18, 2009 06:09 AM

Hey Cole,

Great post, man. I don't have anything to add but I really enjoyed reading your thoughts.

Dave
DNS Reptiles

Sunherp Feb 18, 2009 10:48 AM

Thanks,Dave! I enjoy writing these sorts of things out and working them over in my head. I could very well be on the wrong track, here, but it makes sense to me (and from what I've seen!).

-Cole

cn013 Feb 18, 2009 02:33 PM

Hmmmn well Cole... "temporalis" as a product of a secondary intergrade zone wholistically I'd have to disagree with... lol. Aside from that nice instigatory post! I feel more data concerning the actual migration of the varied subspecies to their currently occupied habitats lends some nice perspective as well. Factor in some cool avg. global temperature shifts and we may find the notion of a truly 'pure' primary intergrade zone to be a rarity. Imagine tossing a few pebbles on some calm water and watching the concentric rings meet and overlap. I'd imagine Shannon is indeed right.... aside from locality all we have are NA milks from varied places.

Where do you mix in taylori in this scheme? Or even celaenops now... I'm sure this will get struck up a bit next time we talk as I'm already finding my expression limited by my slow 'two fingered' typing technique! Anyhow here are three 'temporalis' that may in fact have three distinctly different evolutionary processes...

snake_bit Feb 18, 2009 03:01 PM

Chris when did you get the St Marys female? How old is she?
That Jersey critter is outstanding do you have more pics?
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"Wake me when its April"

Doug L

Sunherp Feb 18, 2009 03:39 PM

Hmmmn well Cole... "temporalis" as a product of a secondary intergrade zone wholistically I'd have to disagree with... lol. Aside from that nice instigatory post! I feel more data concerning the actual migration of the varied subspecies to their currently occupied habitats lends some nice perspective as well. Factor in some cool avg. global temperature shifts and we may find the notion of a truly 'pure' primary intergrade zone to be a rarity. Imagine tossing a few pebbles on some calm water and watching the concentric rings meet and overlap. I'd imagine Shannon is indeed right.... aside from locality all we have are NA milks from varied places.

Where do you mix in taylori in this scheme? Or even celaenops now... I'm sure this will get struck up a bit next time we talk as I'm already finding my expression limited by my slow 'two fingered' typing technique! Anyhow here are three 'temporalis' that may in fact have three distinctly different evolutionary processes...

Chris! This is exactly what I was hoping for!

Let me start at the end of your post and work backwards...

I think we may be seeing 2-3 different processes going on with what we’re calling collectively “temporalis” at the moment, too. Some of the animals are, from what I hear, genetically indistinguishable from syspila and nominate triangulum. These animals, as we know, also LOOK a lot like syspila and are found in the northern portion of “temporalis” range. Other animals, especially the ones from further south, are genetically indistinguishable from elapsoides (which doesn’t seem to cluster with any other milks in genetic studies… suggesting it’s NOT part of the triangulum group). Now, assuming that the Northern and Southern populations actually interbreed somewhere along the coastal plain, we’d have either secondary intergradation (assuming elapsoides is a triangulum that has been isolated by past Climatological events) or introgression (if elapsoides is no longer part of the triangulum group).

So, to sum it up, we may have “temporalis” that are really derived from syspila and nominate triangulum, “temporalis” that are derived from elapsoides, and “temporalis” that are a mixture of the two. I’m assuming you’ve read the same compelling arguments recently that I have… LOL

OK, on to the next topic… primary vs. secondary intergradation!

I don’t find much compelling evidence to suggest that the western milks (taylori, gentilis, multistrata, celaenops, and annulata) are currently, or ever have been, separated from one another in any way enabling them to evolve along separate trajectories. Genetic material has rendered them indistinguishable, and there is current gene flow between all of these adjacent populations. For instance, suitable milk habitat is found in south-western WY, adjacent to UT. Milks have been sighted in that area, though I’m unaware of any vouchered specimens. This suitable habitat is continuous to the south to known taylori haunts, and north, to known multistrata haunts. We find similar situations to the south, were more and more AZ specimens are turning up.

And this brings us to the last point!

It’s my personal feeling that taylori is physically indistinguishable (“average” number of bands is, honestly, an outdated and useless character for identification) from celaenops, gentilis and multistrata without locality data. Taking another step, multistrata and gentilis are indistinguishable from one another, as are celaenops, annulata and amaura. A “subspecies” without discrete diagnostic characteristics (physical or biochemical) is rendered invalid under standard nomenclatural practices. All of the above are, necessarily, subject to the rules of priority and sunk into gentilis (I believe it’s the first named, anyway).

Now, a qualifier: I find the above subspecies designations useful in providing a reference to geographic origin and “typical” appearance of a given animal. Since this is the basic purpose of a subspecies designation, should they really be dropped? I don’t know… LOL

How does syspila fit into this? Well, I’d reason that syspila and nominate triangulum became isolated from the western milks (here-in gentilis) during one or more of the previous glaciated periods on the North American Continent, coming back into contact with one another in the interglacial period(s). This has resulted in the differences we now see between “eastern” milks (syspila and nominate triangulum) and the western forms regarding morphology and ecology in addition to genetics.

-Cole
Image

Deepsea Feb 19, 2009 11:28 AM

Cole is a dork ......... oh snap

jyohe Feb 20, 2009 12:28 PM

the female St Marry's on bottom photo looks like mine more or less....she has grown orange bands instead of white...I hate that...I want white St Mary milks....(so now I gotta get more and hope...LOL...and keep kids )....my male is dark red and albut white yet...he is cool....real deep, dark red....

.....I also noticed last night that the temporalis head is alot smaller / shorter than the eastern milk I have here....small head and face area....cool....but different...

....
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..JY

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Tony D Feb 18, 2009 03:02 PM

Cool post. Don't know too much (anything) about the interface between these two forms but your thoughts were presented well and made sense.
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“Nothing is at last sacred but the integrity of your own mind.” Emmerson

Sunherp Feb 18, 2009 03:40 PM

Thanks, man! How about some photos to keep us satisfied?

-Cole

Tony D Feb 18, 2009 03:53 PM

Been kind of busy moving the collection out but once the dust clears I'll take some more pics. Also been looking at how to house the few critters that will remain. I'm considering using 2 Xterra (sp?) cages that are 36" wide 18" deep and 18" high. They seem to provide good access and visibility and are relatively inexpensive.
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“Nothing is at last sacred but the integrity of your own mind.” Emmerson

tricolorbrian Feb 19, 2009 05:55 PM

I just have one or two things to add, Pales are found on both sides of the Missouri River in SD (if you can even call them Pales, they look more like gentilis than anything). The range for pales in most of SD and Montana is strange in that the average milk snake from there looks nothing like the classic Pales found in N. NE and S. SD. Why is this? Gene flow all the way to MT? Maybe...or maybe just regional variations of gentilis based on soil type and habitat.

Typical screamer Pale from NE

Pale from central SD on the Missouri

I call the bottom snake a gentilis (an UGLY gentilis).

Brian Hubbs, Lampropeltis GOD

tricolorbrian Feb 19, 2009 06:00 PM

Now, for those who think syspila need little a whole bunch of side blotches to be pure:

Syspila from central MO

Pale from northern NE (Oops, look at the red head, of course, Pales can have that too)

tricolorbrian Feb 19, 2009 06:02 PM

BTW, all 4 of the snakes I pictured were found in the wild by myself. Ahem...but, I still have no idea how to classify Pales. It's a conundrum.

Sunherp Feb 19, 2009 08:19 PM

You know how to classify them... don't be silly.

Was that SD animal from Lyman Co.? I can't remember. That animal in the first photo always gets me going. Man, Hubbsy, that beast is sweet. The red-head isn't half bad, either. I'll call you to chat about it.

-Cole

snake_bit Feb 19, 2009 07:24 PM

Brian how rare are central MO milks w/o side blotches ?
Have you seen or found others?
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"Wake me when its April"

Doug L

tricolorbrian Feb 19, 2009 11:56 PM

I have found several w/o side (lower lateral) blotches, or with teeny, tiny ones, but they are not as common as those with lower lateral blotches.

snake_bit Feb 19, 2009 07:32 PM

What would you call this KS milk? And don't say ugly !

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"Wake me when its April"

Doug L

tricolorbrian Feb 19, 2009 11:29 PM

I'd call it a gentilis that looks like a Pale, hence the reason I doubt the Pale is really valid.

terryd Feb 20, 2009 12:24 AM

I'd agree w/ calling multistrata, gentilis, grudgingly.

I like the way things are separated now, if someone tells me they found a taylori it gives me an idea of the area it was found in, and what it should look like.
If all NA milks are lumped as gentilis, then I have to start asking alot more questions to find out where the said gentilis came from. "Oh, Diamond fork canyon, UT., we used to call those taylori."

-Dell

jyohe Feb 20, 2009 12:42 PM

looks pale...but when I looked at the photos properties it says syspila in it....????

..........
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..JY

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terryd Feb 20, 2009 01:34 PM

Jeff,
It is a multistrata x gentilis from CO.

-Dell

The photo properties on this photo say hyposyspila but..........

Image

tricolorbrian Feb 20, 2009 05:44 PM

Yeah Dell, so why are you calling a CO gentilis a syspila? Actually, I don't believe those are intergrades at all. Just gentilis in Weld County. and I'm not saying that because multistrata really doesn't exist, but just because I see no reason to call them intergrades.

Weld County gentilis:

terryd Feb 20, 2009 07:01 PM

All that is is a file property w/ 80 or more other photos.

I don't have a problem calling them gentilis in CO., even if they are in an intergrade zone. So gentilis it is. But they are the most multistrata looking gentilis I've seen. Except yours, that looks like pure gentilils.

Opps gotta go, the wife's yelling at me.

-Dell

snake_bit Feb 20, 2009 07:30 PM

Now thats a cool abberrant gentilis x multistrata .
Which end do you put the pinky in
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"Wake me when its April"

Doug L

jyohe Feb 20, 2009 07:41 PM

spotted integrade with 4 way cross...striped and spotted...see...if you cross 4 subspecies they ahve poorly developed bone structure and look all puffy, no bones means they smoosh all up and look almost like some wierd kind of caterpillar....

sweet larval milksnake..........!

....love bug shots.....
....and does anyone notice some of the fungus they walk by while in the field????....I studied with a mycology club /group for a couple years....wow the things I found when you look around for them.....the purple stuff is cool....and some odd growth forms ...fungus is cool....if anyone wants to see any pics go to the fungus of the month page under Tom Volk's website...I forget what college he works at.?......it should still be valid site....

....good luck......
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......

..JY

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tricolorbrian Feb 20, 2009 08:43 PM

You know...women say I'm a fungi to be with.

Sunherp Feb 19, 2009 07:51 PM

I was waiting for this response! I thought I might be able to call you out of your den with some pushy pale talk. Did you get my message?

We're on the same page (as you know) with the multistrata/gentilis thing. Of course they can have red heads! I hatched one this year like that. Pretty awesome stuff.

RE: syspila - define it! What IS the red milk?

-Cole

Sunherp Feb 19, 2009 07:59 PM

Check out this post above.

Get that getula book out already.

-Cole

tricolorbrian Feb 19, 2009 11:39 PM

A syspila is a Red Milk. A Red Milk is very different from a gentilis, celaenops, multistrata, or taylori. It is more closely related morphologically to amaura and temporalis and Easterns. It generally has those little side blotches or remnants thereof, but not always. It usually has red on the head, wider saddles, and fewer annuli [light rings] (which goes along with wider saddles). True temporalis are actually (my belief) stranded syspila that were cut off by the last ice age. That's all I'm going to say about that...

Sunherp Feb 20, 2009 10:07 AM

LOL Your definition of syspila is the same as mine. Good job!

Now... let's discuss the SD animals! Apparently, we'll have to to it on-line, since you don't know how to answer your phone. I don't think you've posted photos of anything from east of the Missouri. Am I wrong?

-Cole

tricolorbrian Feb 20, 2009 11:16 AM

You are not wrong. And I was out last night when you called. Try calling around 10 AM, or 3 PM, but not after 7PM, except tonite, around 7 should be OK tonite.

terryd Feb 20, 2009 12:03 AM

Tricolor wrote:
"most of SD and Montana is strange in that the average milk snake from there looks nothing like the classic Pales found in N. NE and S. SD."

Hmm, After only seeing a few South Dakota Pales I'd have to agree w/ you, even Walt Deptula stated as much about South Dakota Pales years ago. "South Dakota animals, particularly those from the Black Hills territory, almost appear to be a different subspecies."
I think it will be interesting to see more of these South Dakota multistrata over the next few years to try and get a better handle on what is going on w/ them as they are quiet diffrent looking animals.

Now Montana Pales I'd have to disagree w/ you Brian. There are plenty of classic looking Pales found here that will rival any found in NE. You just haven't seen enough yet.

All field found Montana multistrata.



-Dell

Image

tricolorbrian Feb 20, 2009 12:21 AM

Why don't you show some of the ones you showed me at your house? LOL And, young and sub-adults always look better than adults.
Here's a juvenile central SD snake:

Oh look, here's one of your own MT Pales:

Wow, what a screamer...LOL again...

tricolorbrian Feb 20, 2009 12:27 AM

Here's the type specimen of multistrata (from MT, circa 1855). Yes, it does look more classic.

terryd Feb 20, 2009 12:51 AM

I thought you might say that B man. So here you go.

Photos:
#1. Subadult
#2. Adult
#3. same Pale in #2
#4. Adult
#5. 2 or 3 year old
#6. neonate
#7. 2 year old most likely
#8. Adult

I will agree alot of the Pales in the southern part of the state seem to be a little more newsprinty, and yellow/amber look to them. But that's not a bad thing.
Newsprint doesn't bother me(much), and the yellow/amber animals I kind of prefer.

Adult

Adult

Adult

Adult

Four year old from the male you posted.

Neonates from a w/c female, the egg in the photo is what gives it away.

Image

tricolorbrian Feb 20, 2009 12:57 AM

Dell: I agree there is a lot of variation, and some do look like typical NE Pales, but many do not, hence the need to possibly reclassify them as gentilis (or some new subspecies that looks a lot like gentilis, LOL).

terryd Feb 20, 2009 01:09 AM

Douh! Gentilis. grrr

Syspila adult. aaah, not gentilis.

Image

Sunherp Feb 20, 2009 10:45 AM

That's how we roll, Dell! Nice stuff.

-Cole

jyohe Feb 20, 2009 12:38 PM

the more I look at these "western" milks.....the more they look the same.....makes me wonder....no not all the same...but people post a pic of let's say a celaenops....yet it looks more like an anulata or maybe even a gentilis to me......or pales, all from one area...half look like pales...then one looks like it has celaenops in it...the next might look like a gentils mix etc etc...extra wide white or extra wide red bands can make the snake look like what we see in a different sub-species...

you all get it....just talking......and it actually makes the "locality matters" thing.......actually matter....

.....
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..JY

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tricolorbrian Feb 20, 2009 05:47 PM

You're just confused...anyone can tell the difference between western milks. There is celaenops/taylori and gentilis/multistrata. It's simple when you combine two of the subspecies with adjacent subspecies. LOL Just use the snouts as a reference point. c/t=darker snouts, g/m=lighter snouts.

jyohe Feb 20, 2009 07:46 PM

...I know enough about milks...to get along...but the pics don't all match up with common colors/patterns ALL the time...as you and I can see by the talk in this thread alone....integrade? or not or off color or off patterned....alot of westerns are tough calls without paperwork.....eastern milks are all different but all look like eastern or red or coastal USUALLY....etc etc....

.......SA and CA are a different problem, then add morphs to it....ugh.....

...........this is all why we can all end up with 20000000 snakes if we had room ,money and time.....!....

.
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......

..JY

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Tony D Feb 24, 2009 12:29 PM

Removed from the context of their habitat and their geographic relationship to one another difference can be subtle. Then you read posts like the one Nathen did and you gain a whole new appreciation of the relationship of these animals to their respective habitats.
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“Nothing is at last sacred but the integrity of your own mind.” Emmerson

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