Reptile & Amphibian Forums

True subspecies or?

2.3 Black kings
3.0 hypo Brooksi
0.1 Desert king
1.1 Speckled kings, Louisiana
1.1 Eastern kings, North Carolina
1.1 Lantry Grey bands
1.0 Thayeri buckskin
0.1 Sinaloan
0.1 banded coastal cal king
1.0 aberrant desert cal king
0.1 desert banded cal king
0.1 aberrant sonoran bullsnake, WC

not a clue bud

I guess the "experts" say no. They're pretty cool no matter what.

Todd

forum, I would have to say YES. Subspecification is marginally delineated and intergradation washes out some of its meaning. But as keepers we keep them seperated, so I think it can safely be called an "artificial" subspecies. Just my opinion,Jeff

...strictly from a hobbyist perspective,i say sure why not;absolutely even.....from a taxonomic point of view,well here's where it gets kind of murky.i and many others have found mainland getula in many places that strongly resemble obx kings;lots of stippling/speckling on the dark saddle scales(between the 'chainlinks')....however i'd still call those aberrant/anamolies.hey so what,much of snake taxonomy is what i consider an outdated wreck,with an overhaul on it's way...as time goes on i continue to fall more squarely into the school of 'lumpers' (vs.'splitters')as a general rule(of course there are exceptions)....imho i think anywhere subsp. integrate,sink them into one;where adjacent subsp. don't,well they should probably be elevated to specific status....i.e. all mainland getula subsp. should be sunken,all just being locale-specific morphological variations on a theme......by current definition,seeing as how the kings on the outer banks are in fact isolated genetically from those on the mainland,wouldn't that warrant specific status?
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"with head raised regally,and gazing at me with lidless eyes,he seemed to question with flicks of his long forked tongue my right to trespass on his territory" Carl Kauffeld

I tend to agree that they are a valid subspecies and should be noted for, but I have a feeling this war will rage on for many moons to come!

Very well put! The system that has been in use is longer then we have been alive. Major time for an over haul. If not we might as well go back to calling triangulum, doliata!

imho i'd say the system itself is not the problem at all;the 'overhaul' i was reffering to was the gradual move away from taxonomic inconsistencies,results of somewhat outdated methods-notably,heavy reliance on morphology-with the onset/widespread application of mtDNA....btw watch out,those obx kings have STRONG feeding responses.to feed them all they want/can eat would leed to quite an overstuffed,sausage-like snake,lol.....here's one cb '04 female;hatteras island origin

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"with head raised regally,and gazing at me with lidless eyes,he seemed to question with flicks of his long forked tongue my right to trespass on his territory" Carl Kauffeld

>>True subspecies or?

Ken Krysko, in his work for "meansi" did MtDNA analysis on the Outer Banks Kings and found them to be genetically identical to mainland forms as far away as S. Georgia (L.g.getula). Given this, it's hard to argue them as being a seperate ssp. More a pattern variant like the striped and banded Cali Kings of the west.
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Ace

Ken Kyrsco got a $1000,000.00 grant and did his thesis on the eastern getula. DNA does not lie.
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"Yeah ya told me, and ya wrote it down too. But how the hell am I supposed to remember!"

Give me a $1,000,000 grant and I will PROVE water isnt wet, the sky isnt blue, and snakes are just hiding their feet!LOL

Give me a $1,000,000 grant and I will PROVE water isnt wet, the sky isnt blue, and snakes are just hiding their feet!LOL

I get what you are saying but Krysco was not trying to prove anything with the OB's. If anything the results from the goini were more interesting as well as the range of floridana and the easterns kings meet.
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"Yeah ya told me, and ya wrote it down too. But how the hell am I supposed to remember!"

What did the results say about goini?
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Mike
KingPin Reptiles Inc.
www.kingpinreptiles.com
^ Updated 7/15

According to the paper, goini is not valid and is a relict population of getula x floridana (no surprise there); however, they did find support for a ssp that is patternless or near patternless and is fairly isolated in a certain portion of the habitat ("meansi" after Bruce Means, the co-author of the paper).

Isnt that what he WANTED to prove going in? Isnt Means more "familiar" with those than any other "ssp"? Coincidence? I have to disagree. There is the skeptic in me thinking he wants to sink "goini" and put his own nomicer on it, which makes the ssp validations a little more suspect than what Williams did in his findings about "coastal plains". If someone can point me to the SCIENCE I would appreciate it,thanks,Jeff

He puts no ssp name on the locality snakes but points out the largely phenotypical characters and mtDNA sequencing.

This paper was 2001 and largely discounts goini, sticticeps and brooksi using rather sound arguments.

Chris

What is the papers title? I'd like to read it.

Thanks
Vinny
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“There is a grandeur in this view of life, with its several powers, having been originally breathed into a few forms or into one; and that whilst this planet has gone on cycling according to the fixed laws of gravity, from so simple a beginning endless forms most beautiful and most wonderful have been, and are being, evolved.” -C. Darwin, 1859

Natural Selection Reptiles

ECOLOGY, CONSERVATION, AND MORPHOLOGICAL AND MOLECULAR
SYSTEMATICS OF THE KINGSNAKE, Lampropeltis getula (SERPENTES:
COLUBRIDAE)
By
KENNETH L. KRYSKO

A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL
OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT
OF THE REQUIREMENTS FOR THE DEGREE OF
DOCTOR OF PHILOSOPHY
UNIVERSITY OF FLORIDA
2001

Chris

Here's a piece of his paper reproduced WITHOUT permission (hope he doesn't mind)

Phylogeography
All morphological and molecular analyses performed in this study support a western
origin of L. getula in the eastern U.S. The evolutionary history of eastern L. getula
appears to be linked to the geological history of the Coastal Plains derived from ancient
coastlines. Fossil remains indicate that ancestral western L. getula utilized a Gulf Coast
corridor to disperse east into Florida during low sea levels in the Pliocene between 2-5
million years ago (Mya) (Auffenburg, 1963). Estimates of mtDNA sequence divergence
for reptile species range from 0.47 to 1.32% per million years (Zamudio and Greene,
1997). The smallest percentage sequence divergence between the eastern and western
clades of L. getula is 5.56%, which translates into 4.2-11.8 Mya, a time frame that
overlaps the sequence of events hypothesized by Auffenburg (1963). Although some
midwestern samples were much closer geographically to eastern samples, they are much
less closely related relative to the range of divergences within the eastern clade. A rapid
range expansion from southern refugia into northern areas as glaciers retreated could
account for the high haplotype diversity and low genetic distances (maximum d = 0.91%)
in the east in comparison with other geographic areas. Interestingly, additional snake
taxa with similar distributions in the midwestern and eastern U.S. have been reported to
exhibit similar phylogeographical patterns (Rodriguez- Robles and Jesus- Escobar, 1999b,
1999a; Burbrink et al., 2000; Burbrink, 2001). Thus, the evolution of the three identified
123
subclades within the eastern clade is probably related to more recent glacial events during
the Pleistocene (10,000-2 Mya).
The lowest sequence divergence between the eastern and midwestern clades is
2.18%, which translates into 1.65-4.6 Mya (early Pleistocene to early Pliocene). There
were four major glacial events during the Pleistocene, with the first being the greatest
followed by less intense episodes (Webb, 1990; Brown and Lomolino, 1998). During
these glacial events sea level was >100 meters (m) lower than present-day (Lidz and
Shinn, 1991), Florida was nearly twice its present size and land was continuous from the
present Florida mainland to the Dry Tortugas (MacNeil, 1950; Hoffmeister and Multer,
1968). In order to escape from cooler climates and ice masses, extreme northeastern L.
getula populations must have been extirpated or pushed southward, whereas peninsular
and panhandle populations experienced an expansion following suitable habitat. These
conditions of displacement and expansion would have promoted mixing among SE U.S.
populations. This is evident because of their close genetic relationships as well as certain
shared morphological characters such as ontogenetic interband lightening found in
Florida and as far north as the Outer Banks of North Carolina (Barbour and Engles, 1942;
Lazell and Musick, 1973; Blaney, 1977). However, a less likely but alternative
hypothesis is that the character of ontogenetic interband lightening evolved multiple
times along the Coastal Plains. During major interglacial events much of present-day
Florida was inundated except for higher elevations in the peninsula and panhandle.
These areas of higher elevations have been identified as ancient shorelines in the
peninsula (Jackson, 1973; Webb, 1990; Clark et al., 1999), as well as ancient barrier
islands in the eastern Apalachicola Lowlands (Brenneman, 1957; Brenneman and Tanner,
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1958). During these times, Atlantic populations were capable of expanding their range
northward following warmer climate, but peninsular and panhandle populations were
geographically isolated. Peninsular populations were probably found in mesic habitats
extending into the southern peninsula (Watts and Hansen, 1988). Panhandle populations
were probably isolated on barrier islands in the eastern Apalachicola Lowlands
(Brenneman, 1957), which correlate well with a large number of endemic and relict
plants and animals there (Table 3-10; James, 1961; Clewell, 1977; Yerger, 1977; Ward,
1979; Judd, 1982; Gilbert, 1987; Coile, 1996; Chafin, 2000; Chaplin et al., 2000). Thus,
the distributions of the Apalachicola Lowlands endemic plants and animals possibly were
a result of the same geological event. Since the last glacial maximum around 18,000
years before present (BP), North American glaciers have melted and retreated northward,
sea level has risen steadily (Randazzo and Halley, 1997). Again, populations of eastern
L. getula will eventually become isolated in the peninsula and panhandle, while others
expand their range northward.
Taxonomy
Under the Apomorphic Species Concept (ASC), species are considered to be wellsupported
minimal monophyletic groups. In all morphological and molecular analyses in
this study, the eastern populations of L. getula represent the smallest well-supported
monophyletic group, which suggests recognition as a separate species. Recently, users of
the various Phylogenetic and Evolutionary Species Concepts (PSC and ESC respectively)
have objected to the subspecies concept because they claim it is not operational within
their definitions of reciprocal monophyly (Mayr and Ashlock, 1991; Baum, 1992;
Burbrink et al., 2000). Additionally, beginning with Wilson and Brown (1953) the
125
subspecies concept has been criticized because subspecies were commonly identified on
too few and arbitrary delimited characters. And in many cases where several characters
were used, each character varied independently because of differing locally adaptive
pressures, which resulted in different subspecies distributions and arbitrary cline slices
(Frost and Hillis, 1990). Nonetheless, under the ASC one may recognize subspecies as
subclades within species, where subclades have weaker support because of occasional
gene flow or recent evolutionary divergence (Mishler and Theriot, 2000). Mayr (1969)
and Smith et al. (1997) also recognize subspecies as geographic races with relatively
homogenous phenotypic patterns, which have the ability to exchange genes with adjacent
races. Areas where subspecies come into genetic contact are identified as intergradation
zones (Mayr, 1969; Smith et al., 1997). Because gene flow will obscure boundaries of
subspecies and prevent them from attaining reciprocal monophyly at the mtDNA level,
this criterion should not invalidate subspecies recognition (Patton and Smith, 1994;
Rodriguez-Robles et al., 1999c).
The traditional subspecies recognized in L. getula in the eastern U.S. correspond
closely to the phylogenetic patterns that I have uncovered. After examination of
hundreds more specimens than any previous researcher for a study of geographic
variation using morphology (Chapter 3), I identified three subspecies in Florida: 1) L. g.
floridana from central and southern peninsular Florida, 2) L. g. getula from NW
peninsular Florida north to southern New Jersey, and 3) unnamed populations in the
eastern Apalachicola Lowlands. In the morphological and molecular analyses, the three
consistently identified subclades correspond exceptionally well to the three identified
geographic races (Chapter 3). Because each of these subspecies can be diagnosed by at
126
least one synapomorphy, this evidence suggests that they represent evolutionary
processes rather than ecotypic variants.
Individuals from the western Apalachicola Lowlands were once believed to
represent a distinct subspecies, L. g. goini (Neill and Allen, 1949). However, Blaney
(1977) invalidated L. g. goini by speculating that it represented a Pleistocene intergrade
between panhandle L. g. getula and now disjunct peninsular L. g. floridana.
Additionally, Means (1977) believed that L. g. goini indeed represented an intergrade
population, but L. g. goini was intermediate between unnamed populations in the eastern
Apalachicola Lowlands and L. g. getula that surrounds the region. The most consistent
subclade with statistical support in morphological and molecular analyses consisted
primarily of eastern Apalachicola Lowlands and panhandle individuals. Additionally,
there are five synapomorphies supporting the monophyly of the eastern Apalachicola
Lowlands populations. Morphological data examined on a much finer geographic scale
(Chapter 3) revealed that the eastern Apalachicola Lowlands populations possess
relatively homogeneous color patterns that grade into populations in surrounding areas
(i.e., gene flow) consisting of L. g. getula. Because L. g. goini possesses intermediate
characters between eastern Apalachicola Lowlands snakes and L. g. getula, I relegate L.
g. goini to intergrade status. Therefore, because L. g. getula and L. g. floridana are
already recognized subspecies, the eastern Apalachicola Lowlands populations deserve
equivalent taxonomic recognition as well. As systematists it is our responsibility to
document biological diversity, and it is essential to recognize both species and subspecies
otherwise everyone worldwide will fail to notice biological diversity without names
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(Dobzhansky, 1970; Smith et al., 1997). A taxonomic amendment regarding the eastern
Apalachicola Lowlands populations is in progress and will be treated in a separate paper.
I was unable to find evidence supporting the recognition of both L. g. brooksi and L.
g. sticticeps using morphology and mtDNA. Individuals from the extreme southern
Florida peninsula were once believed to represent a distinct subspecies, L. g. brooksi
(Barbour, 1919), yet its validity quickly came into question (Blanchard, 1920, 1921;
Wright, 1935). Becasue only 15 individuals from the Florida peninsula were sequenced
for mtDNA, this small sample size might have been insuffiecient to find genetic
partitioning within the peninsula. Nonetheless, Duellman and Schwartz (1958), Blaney
(1977), Krysko (1995), and Krysko (this study) could not find morphological characters
for diagnosing L. g. brooksi, and the mtDNA data indicate that the Florida peninsula is a
panmictic population (Tables 5-4, 5-5). Barbour and Engles (1942) described L. g.
sticticeps from the Outer Banks of North Carolina based on head morphology and
ontogenetic interband lightening of the dorsal pattern (Lazell and Musick, 1973).
However, Hillestad et al. (1975), Blaney (1977), Gibbons and Coker (1978) and Palmer
and Braswell (1995) did not accept that a distinct geographic race existed on the Outer
Banks. My morphological and molecular data further corroborate these authors, as I
found snakes from the Outer Banks with mtDNA haplotypes identical to adjacent
mainland snakes as well as those as far away as SW Georgia.

2006 - Deals mostly with "Meansi"

http://www.brucemeans.com/pdf/2006KryskoandJuddLampropelt.pdf

2001 - Deals mostly with Floridana

http://purl.fcla.edu/fcla/etd/UFE0000326

If he plays by the rules and goini is found to be invlaid then he cant give it a new name, it becomes synonymous with L. getula. If other work finds that goini is infact a valid ssp. then it cant get a new name the old name goini has to be resurrected. Ah, the rules of the ICZN

Vinny
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“There is a grandeur in this view of life, with its several powers, having been originally breathed into a few forms or into one; and that whilst this planet has gone on cycling according to the fixed laws of gravity, from so simple a beginning endless forms most beautiful and most wonderful have been, and are being, evolved.” -C. Darwin, 1859

Natural Selection Reptiles

>>If he plays by the rules and goini is found to be invlaid then he cant give it a new name, it becomes synonymous with L. getula. If other work finds that goini is infact a valid ssp. then it cant get a new name the old name goini has to be resurrected. Ah, the rules of the ICZN

Intersting that you mention this. In Krysko's paper that he co-authored with Judd (2006) he describes what he considers what a true meansi is, and the pattern variations of meansi. ALL the goini holotypes and paratypes fall within his range of variation of meansi, making meansi nothing more than a rediscription of goini.

Now this part confuses me...... He claims goini is an intergrade of meansi and L.g.g., but finishes by including them as a pattern variant. So, according to ICZN would this also make meansi an intergrade (with??) since he also considers them to be pattern variants of the same snake (goini), or does it make goini valid again?

Personally, I think all the Apalachicola Kings are pattern variants of L.g.g., and Kevin Enge's work with the Mosiac Easterns shows exactly how these variations came about.
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Ace

It shows that with a budget, a bunch of educated words, cited work twisted into what you are looking for,backup from a guys with a PHD after their name, you can validate or invalidate pretty much whatever you want. I am going to reread the paper in detail again, but there are more than a few of those "inaccuracies". Reminds me of a golfer picking up his lie, rubbing the ball, pretending to clear away debris, and returning the ball to the exact same spot. For someone watching, it makes NO SENSE to have spent money,time,etc. All the "findings" are based upon hypothesese that cant be "proven". I wont worry,I eat crow with the best of them,lol. After all, with the known variances in pattern, why do we bother with subspecification? You cant protect habitat based upon a pattern variation.That just wont hold up in court......

When and where was "goini" ever considered valid?

It's been expressed in quotes as long as I've been into snakes.

Chris

"Goini" was determined to be the result of easterns intergrading with meansi (not floridana).

Meansi was determined to be patternless, striped and banded kingsnakes within a relatively small range (a section of 2 counties). Patternless and striped kings are still found in the intergrade/"goini" range, and shouldn't be considered the mark of a true meansi.

...what you call 'em anymore but some of my eggs are starting to pip. I think I'm callin' 'em Chipola R. kings
Here's one of my females....

:Mark
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Surrender Dorothy!

www.crimsonking.funtigo.com

I wonder what Brian Hubbs is going to say in his book about these.
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"Yeah ya told me, and ya wrote it down too. But how the hell am I supposed to remember!"

They are PURE Awesome!

and that one is a beauty... be sure to send me pix once they emerge!

~Z
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Royal ReptileZ

~ZF
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Royal ReptileZ

not quite how i read it;what neill and allen found('goini') in 1949 was ultimately determined to be an integrade between what is now 'meansi'(patternless and/or striped individuals only);any presence of bands is indicitave of l.g. getula influence from the surrounding north/east/west.....so to bring this back to the topic-blaney(whom krysko says to concur with)sunk sticticeps,considering obx kings as simply a relict intergrade of floridana and getula-meanwhile krysko finds them genetically identical to kings from the adjacent mainland of n.c. and even georgia...i would like to know why,if an geographically isolated pop.(sticticeps) is not valid,how can you not sink the confluent splendida,holbrooki,nigra,et al.?...where is the mtdna or satelite dna work with these other mainland froms that all bleed together?....also noticed krysko mentions 'ancestral western getula'....a side note- now maybe i'm not thinking clearly-but if a taxonomic reconfiguration were to happen then i'm guessing the nominate(or specific title,if all subsp. are wiped out) would be californiae(or nigrita?).....btw gorgeous kingsnake,mark
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"with head raised regally,and gazing at me with lidless eyes,he seemed to question with flicks of his long forked tongue my right to trespass on his territory" Carl Kauffeld

So where does this leave the outer bank king, as a mixed, mongrel hybrid?

>>i would like to know why,if an geographically isolated pop.(sticticeps) is not valid,how can you not sink the confluent splendida,holbrooki,nigra,et al.?

Apparently stricticeps hasn't been isolated long enough to form it's own genetic variation. It only stands to reason that different coast species will have greater genetic variation than same coast variations. Afterall, they've been seperated much longer than same coast forms.

>>>...where is the mtdna or satelite dna work with these other mainland froms that all bleed together?

Krysko used outgroups of nigra, splendida, and californiae (If I remember correctly). He found that nigra showed more than 2% MtDNA variation from the Eastern forms, and the western forms showed over 5%.

>>>....also noticed krysko mentions 'ancestral western getula'....

The ancestral western form is assumed to be splendida. Many believe splendida (or something similar) to be the founding stock to all the present getula forms. I believe Blanchard was the first to introduce this idea in his work "A Revision of the Kingsnakes" (1921).

>>>a side note- now maybe i'm not thinking clearly-but if a taxonomic reconfiguration were to happen then i'm guessing the nominate(or specific title,if all subsp. are wiped out) would be californiae(or nigrita?)

Actually, it'd be getula (first form described). Most likely being split into three seperate species/groupings with no ssp. The Eastern clade (L.getula) consisting of L.g.g., meansi/goini and stricticeps, the central clade (L.holbrooki) consisting of holbrooki and nigra, and the western clad (L.californiae) consisting of californiae, splendida, and nigrita. Floridana seems to be at odds with the Eastern clade in having higher dorsal scale counts, but similar hemipenile shape, so I'm not sure where they'd go.
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Ace

...i mentioned;i saw that,how krysko did indeed sample some of the midwest/western forms.but his focus was the eastern clade-as far as i know no such detailed work has been published re: the others.will be interesting to see the taxonomic 'aftermath' of such work.you pointed out genetic variations of up to 5% from coast to coast-and yet there is plentiful evidence of integration between all subspecies involved,clear across the country.the three division approach you mention makes plenty of sense to me,but so would eliminating the subsp. altogether in this case,imho....as for the nominate form point,i couldn't remember if that would revert to the first form described,or the accepted ancestral form.appreciate the informed feedback;interesting thread here.
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"with head raised regally,and gazing at me with lidless eyes,he seemed to question with flicks of his long forked tongue my right to trespass on his territory" Carl Kauffeld